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HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 1

NOTES ON THE AMERICAN SOFT-SHELL TURTLES
WITH SPECIAL REFERENCE TO AM YD A AGASSIZII

By Leoxhard Stejxeger

With Thirty Plates

CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
May, 1944

PUBLICATIONS
OF THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
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Of the Bulletin, Vols. I to XCIII, and Vol. CXIV, No. 1
have appeared and of the Memoirs, Vol. I to LVI

These publications are issued in numbers at irregular intervals.
Each number of the Bulletin and of the Memoirs is sold separately.
A price list of the publications of the Museum will be sent upon
application to the Director of the Museum of Comparative
Zoology, Cambridge, Massachusetts.

After 1941 no more Memoirs are to be published.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 1

NOTES ON THE AMERICAN SOFT-SHELL TURTLES
WITH SPECIAL REFERENCE TO AMYDA AGASSIZII

By Leonhard Stejneger

With Thirty Plates

CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM

May, 1944

PREFATORY NOTE

T. Barbour

As has been well known, for many, many years before his death
Doctor Stejneger was engaged in the preparation of a general treatise
on the fresh water and land Testudinata of North America. After his
death it was found that by far the greatest part of his notes were in the
fragmentary state that often holds in an early stage of preparation
for publication. The only section which might be said to be in pub-
lishable form and to reflect the mature conclusions of the author was
this part comprising his summary of the genus Amyda. For many and
obvious reasons it seems desirable to bring this to public attention and
by the same token it seems presumptuous to change it.

The Museum of Comparative Zoology has the good fortune to
present to the public what will perhaps be the last example of the work
of this distinguished veteran worker in our field. By the consent of the
authorities of the United States National Museum this institution has
permission to publish this work, a tribute of honor and respect to the
author, being published here only because the situation induced by
the state of war makes it inconvenient for the Government Printing
Office to handle the work at this time.

No. 1. — Notes on the American Soft-Shell Turtles ivith Special Reference

to Amy da agassizii1

By Leonhard Stejneger

INTRODUCTION

The number of living indigenous soft-shell turtles of the family
Trionychidae in North America is less than a half dozen, while more
than a half hundred fossil forms, the earliest dating as far back as the
Cretaceous, have been described by paleontologists from the same
region which embraces the habitable part of North America east of
the Rocky Mountains and northern Mexico.

The living American forms have been considered from time to time
by prominent herpetologists as belonging to several genera under
varying names, according to their views on the genotypes. Thus
Agassiz recognized three genera and so did Baur and Hay; Cope and
True reduced the generic subdivisions to two. Boulenger, Siebenrock,
and recent systematists have regarded them as congeneric; the name
has varied accordingly.

NOMENCLATURE

In the early stages of systematic herpetology, before the idea of the
"genotype" had become generally accepted, there was no fixed rule
which would designate the type of genera with more than one species.
A sort of "selection by the first reviser" was early practiced. Thus if
an author subsequent to the creator of a plurispecific genus, in sub-
dividing the latter, retained this name for a single species, applying
one or more new names to the rest of the species, his selection, as a
rule, was respected, and thus the idea of type selection by "elimina-
tion" became a more or less general practice. But because of the un-
willingness or ignorance of some workers and the varying and uncer-
tain application of this method, great confusion in the nomenclature of
genera resulted. This is what happened to Geoffroy's generic term
Trionyx (1809), which has been the cause of so much confusion. In
1816 Oken was the first one to subdivide the plurispecific genus, re-
serving as he did the name Trionyx for the single species T. granosa,
leaving the others under the name Amy da. In 1830 Wagler reported
Oken's monotypic selection of Trionyx granosa, but substituted As-
pidonectes for Amyda. The confusion was caused by Gray who in his

1 Published with the aid of a special gift from Mr. G. R. Agassiz.

6 bulletin: museum of comparative zoology

Synopsis Reptilium; 1831, erected the monotypic genus Emyda for
the same species. He did so in ignorance of Wagler's action and after
seeing Wagler's work called attention to it in the "Additions and Cor-
rections" in the same work, p. 78. Yet in the "Catalogue" of 1844 and
his later writings he still retained Emyda. Gray was followed by
Boulenger and many writers since 1889, in spite of Baur and Hay who
in 1898 and 1904, like Bonaparte in 18361 and 1857'2, upheld Wagler's
action.

Following these dissertations by Baur and Hay, the case of the
genotypes of the genera Amy da and Trionyx was discussed by me in
1905 (Science, new ser., vol. 25, Feb. 10, 1905, pp. 228-229), conse-
quently before the adoption (1908) of the present wording of Article
30 of the International Rules of Zoological Nomenclature. In 1905 the
argument was based on the process of elimination, which at that time
was considered the legitimate process. On that occasion, the type of
Trionyx was decided to have been selected in 1816 by Oken as first
revisor by segregating Trionyx granosus, supposed to be synonymous
with Geoffroy's T. coromandelicus, as the sole species in the restricted
genus.

However, the adoption in 1908 of the changed form of Article 30
apparently nullified previously accepted type selections unless they
were couched in the stringent language of the new version.

The new Article 30 was framed and phrased for the purpose of doing
away with the uncertainties of personal interpretation unavoidable in
the process of elimination. It was intended that the new rule should be
applied literally; hence the first article (I a) reads: "When in the origi-
nal publication of a genus, one of the species is definitely designated as
type, this species shall be accepted as type, regardless of any other con-
siderations." (Italics mine.) To emphasize this intention the last
paragraph of the article was inserted as follows: "The meaning of the
expression 'select the type' is to be rigidly construed. Mention of a
species as an illustration or example of a genus does not constitute a
selection of a type." (Italics mine.)

Applied to the case in hand the facts are these:

In the original publication of Trionyx by Geoffroy in 1809 (Ann.
Mus. Hist. Nat., Paris, 14, pp. 1-20, pis. 1-5) none of the many species
was definitely designated as type. Not in the body of the text proper,

1 Cheloniorum Tabula Analytica, p. 8: "22 AMYDA, Schweigger (Aspidonectes WAGL.
Trionyx GRAY. BELL. Gymnopus, DUM." "23. TRIONYX WAGL. {Emyda, GRAY.
BELL. Cryptopas DUM.)"

2 Contrib, Nat. Hist. United States, 1, p. 330: "Trionyx proper in contradistinction

to Aspidonectes"; see also pp. 394-395.

stejneger: American soft-shell turtles 7

but in the Explication des planches {explanation, of the plates) at the end
of the article, occurs, however, the following mention of the trionyx
of Egypt as an illustration and example of the genus, which was
omitted in the simultaneous reprint of the article in the Bulletin of the
Philomatic Society of Paris :

"Le trionyx d'Egypte, represente planche I, vue en dessous el de cote,
nous donnant une idee exacte du port et des caracteres generiques des
trionyx, nous sommes bornes, dans les planches suivantes, a faire
figurer les seules parties caracteristiques des autres especes, telles que
leurs carapaces en a, et leurs plastrons en B." l

Even if this kind of mention as an "illustration and example" may
be characterized as a "clear intention" it is certainly not a definite desig-
nation rigidly construed.

Consequently, Geoffroy having failed to definitely designate T.
aegyptiacus as the type of Trionyx, it was left to the first subsequent
author to select the type, and that was done by Fitzinger in 1843
(Syst. Rept., p. 30) who selected T. coromandelicus Geoffroy,2 "and
such designation is not subject to change."

Having now disposed of the type designation of Trionyx for a genus3
in the subfamily Cyclanorbinae, the question of the status and type of
Amyda arises.

Among the new species described by Geoffroy in the same paper of
1809, is the Trionyx javanicus (Ann., p. 15). After a brief diagnosis he
added the following svnonvm:

"Amyda javanica SCHNYEIGGER, dans un manuscrit communique
a l'Institut."

This publication of Schweigger's monotypic generic name Amyda
clearly establishes its availability for the species congeneric with his
Amyda javanica, the description of which by Geoffroy is regarded as
identical with Boddaert's Testudo cartilaginea of 1770.

Fitzinger's use (1835) of the name Amyda for a section ( = subgenus)
with Trionyx subplamis as type, and Agassiz's subsequent (1857) ap-
plication of the same name for another genus with Amyda mntica as
type are consequently both void. Dogania is the proper name for the
former; Euamyda is now available for A. mntica as a generic or sub-
generic name.

1 The Egyptian trionyx, represented on pi. 1, viewed from below and from the side, gives us an
exact idea of the aspect and the generic characters of the trionyxes; we limit ourselves in the
following plates to figure only the parts characteristic of the other species, such as their carapaces
in a and their plastrons in b.

2 Trionyx coromandelicus Geoffroy, 1809, is a synonym of Testudo granosa Schoepff, 1792

3 Synonyms: Emyda Gray, 1831 (not of Rafinesque, 1815); Lissemys Malcolm Smith, 1931.

8 bulletin: museum of comparative zoology

Genus Amyda1 Schweigger

1809. Amyda SCHWEIGGER in Geoffroy, Ann. Mus. Hist. Nat., Paris, 14,
p. 1 (monotype Amyda javanicus = T. cartilagineus).

1816. Amyda OKEN, Lehrb. Zool., 2, p. 348 (type designated by Stejneger,
1907, Trionyx euphraticus).

1830. Aspidonectes WAGLER, Nat. Syst. Amphib., p. 134 (type designated

1843 by Fitzinger, Trionyx aegyptiacus = T. triunguis).

1831. Trionyx GRAY, Synops. Rept., p. 45 (type T. ferox) (not of Oken,

1816; Wagner 1830).

1835. Gymnopus DUMERIL and BIBRON, Erpet. Gen., 2, p. 472 (substi-
tute name for Aspidonectes Wagler) (p. 484 Gymnopodus, laps,
calam.)

1835. Platypeltis FITZINGER, Ann. Wien Mus., 1, pp. 120, 127 (type desig-
nated by Fitzinger 1843, Platypeltis ferox.)

1835. Pelodiscus FITZINGER, Ann. Wien Mus., 1, pp. 120, 127 (type desig-
nated by Fitzinger 1843, P. sine?isis.)

1842. Aspedonectes HOLBROOK, North Amer. Herp. 2 Ed, 2, p. 18

(emendation).

1843. Potamochelys FITZINGER, Syst. Rept., p. 30 (monotype Aspidonectes

javanicus).

1844. Tyrse GRAY, Cat. Tort. Brit. Mus., p. 47 (type T. nilotica = T. triun-

guis).

1856. Tryonix SAGER, Peninsular Journ. Medic Coll. Sci., 3, no. 8, Feb. 1856,

p. 361 (emendation).

1857. Amyda AGASSIZ, Contr. Nat. Hist. United States, 1, p. 398 (mono-

type A. mutica) (not of Schweigger).

1864. Rafetus GRAY, Proc. Zool. Soc. London, May 1864, p. 81 (monotype
T. euphraticus).

1864. Aspilus GRAY, Proc. Zool. Soc. London, 1864 (p. 83) (type Aspilus
cariniferus).

1869. Callinia GRAY, Proc. Zool. Soc. London, 1869, p. 221 (type, T.
spiniferus).

1895. Platyrettis KIRSCH, Bull. U. S. Fish Comm. 1894, p. 333; typogr.
err.? for Platypeltis).

1900. Aspidonectus BEYER, Proc. Louisiana Soc. Nat., 1897-1899, p. 43
(emendation).

1 Name of uncertain origin, but apparently a variant of Emys, a river turtle. In the synonymy,
the numerous generic terms based on exclusively Old World species have been omitted. Refer-
ence to these may be found in Bulletin United States National Museum No. 58, 1907, p. 514.

stejneger: American soft-shell turtles 9

The generally accepted five species of the wider genus Amy da in
North America naturally fall into three groups.

1. The A. mutica group, by Agassiz considered a distinct genus and
by many accepted as a subgenus (for which the name Eliamyda
may be substituted as Amy da Agassiz is preoccupied).

2. The A. feroz group including the species A. ferox, spinifera and
emoryi.

3. The A. agassizii group containing only one species on this conti-
nent.

The external appearance of these turtles is so much alike that the
early naturalists had difficulty in diagnosing them properly, with the
result that their taxonomic history is full of misidentifications and mis-
conceptions, even to the extent that the first group, at least on one
occasion, has been suspected of being only the sexual form of one of the
species of the second group.

As in so many of the Old World soft-shelled turtles the essential
characters are recognizable only in the bony structure, so also in our
American species. The most important ones are found in the skull and
the plastral bones and they will form the main subject of the following
discussion.

Skull Characters

The relative proportions of the skulls, their component bones and
the size and shape of the various fossae and foramina vary enormously
with age, hence comparisons have to be made between specimens of
about the same size. The individual variability, which is considerable,
increases with age very often to such a degree, — for instance the
enormous expansion of the alveolar surfaces in the old specimens of
A. ferox, — as to make even group definition difficult. Measurements
of a large number of skulls have therefore to be made, and as skulls of
approximately equal size are rarely to be had, the measurements have
to be reduced to percentages of some standard dimension. As such I
have selected the basicranial length (posterior edge of occipital con-
dyle to tip of snout). The analysis of numerous measurements has
convinced me that skulls with a basicranial length between 40 and 70
mm. (average about 55 mm.) practically represent the normal propor-
tions of important cranial dimensions of our American trionychids.
The relative proportions of the various parts have at that size reached
a sufficient stability and the figures consequently are comparable inter
se. Unfortunately, series of skulls of soft-shelled turtles are not numer-

10 bulletin: museum of comparative zoology

ous in museums, especially when reduced to specimens with a basi-
cranial length between 40 and 70 millimeters, hence the tables pre-
sented below, based as they are exclusively on United States National
Museum material, total only 41 specimens; nevertheless, the figures
are believed to be fairly representative.

An inspection of table 1 will show that Amy da agassizii possesses
smaller internal choanae (ch, plate 1, fig. 2) than the other species, but
a slightly larger intermaxillary foramen (int. max. for., plate 1, fig.
2). Were it not for the latter fact the distance between the inter-
maxillary foramen and the choanae would have been greater than it
is, (viz. 12.5 against only 7.7 to 9.7) in the other species; in other
words, in A. agassizii the longitudinal diameter of the choanae equals
their distance from the intermaxillary foramen, while in the others it
is much greater, in A. ferox even averaging twice as much.

The table further shows the greater width of the alveolar surface of
A. agassizii at the intermediary age of these specimens, but older males
(over 70 mm. basicranial length) of A. ferox acquire an increasing
width of the alveolar surfaces far exceeding in proportion even that of
A. agassizii (plate 30). The width of the alveolar surface of the
mandible in A. agassizii is twice, or nearly twice, as wide as in the
others of the corresponding size.

On the underside of the skull A. agassizii, in addition to the small
size of the choanae and the greater width of the alveolar surface, is
characterized by the position of the suture between the palatines and
the basisphenoid relative to the posterior edge of the temporal fossa.
This is coincident with the different shape of the opening of the tem-
poral fossa, the posterior edge of which is much wider and nearly at a
right angle to the axis of the skull. If therefore a line is drawn across
the base of the skull at the level of this edge, the line passes nearly
through the palatine-basisphenoid suture, while in the other species it
crosses the basisphenoid at or slightly anterior to the middle (plates
1, fig. 1; 6, 30.

The skull of A. mutica is unique among the Amydas in the slender-
ness and delicacy of its bones, especially those of the snout, which is
exceedingly narrow and elongated. The distance from the tip to the
posterior rim of the orbit is much greater than from this point to the
posterior edge of the tympanic cavity, while in the other American
species it is much shorter. This difference is mainly caused by the
reduction in A. mutica of the temporal fossa, the diameter of which
is about six tenths of that of the orbit, while in the other species it
equals or averages even slightly greater than the orbit. The weakness

SI KJNEGER: AMERICAN SOFT-SHELL TURTLES

of the A. mutica skull is further emphasized by the narrowness of the
interorbital spaee and the practical absence of an alveolar surface on
the mandible.

The normal skulls of the remaining American species, A. ferox, A.
spinifera and A. emoryi are essentially alike and present no striking
differences from the normal Amyda skull. They agree with A. agassizii
in the greater massiveness of the bones and general proportions of the
parts. With A. mutica they agree in the longer choanae and the con-
sequent shorter distance of the latter from the intermaxillary foramen.

Table 1

Cranial measurements reduced to
basicranial length of skull

c3
fcC

spinifera

s-
O

ferox group
collectively

'-3

Basicranial length in millimeters,

average

51.8

58.3

53.8

54.0

42.8

Range of b. 1. of specimens

44-70

45-70

46-69

44-69

40-48

Number of specimens measured

Tip of snout to orbit

28.6

24.9

26.0

25.2

25.4

28.71

Orbit, horizontal diameter

19.1

20.0

19.9

20.2

20.0

20.6

Orbit to tympanic cavity

31.7

36.0

27.1

31.5

20.6

Temporal fossa, longest diameter

19.3

23.7

19.3

20.7

21.2

12.6

Interorbital width

8.5

6.0

7.4

7.8

7.1

5.1

Maxillary alveolar surface, width

11.4

6.8

7.4

8.9

7.7

5.1

Internal choanae, length

12.7

15.4

16.2

15.4

15.7

14.7

Choanae to intermaxillarv foramen

12.7

7.7

9.1

8.9

8.6

9.8

Intermaxillary foramen, length

12.2

11.3

11.2

9.8

10.8

15.6

Mandibular symphysis, length

19.5

11.8

13.2

14.3

13.1

20.3

Mandibular alveolar surface, width

10.6

5.5

5.4

5.9

5.6

1 Four specimens only, due to the fact that the bones — intermaxillaries and maxillaries of
this species are so fragile or poorly ossified that they were lost or mutilated in the preparation
of the others. The omission in .4. mutica of a measurement of the alveolar width of the mandible
indicates that it is so slightly indicated that it almost may be said to be nonexisting.

12 bulletin: museum of comparative zoology

The great changes in shape, proportions, color pattern and structure
of the soft -shelled turtles according to sex, age, and individual variabil-
ity make it impracticable to construct a workable key to the species,
hence I have confined myself to specimens in their early maturity,
when the critical characters have assumed a relative and comparable
stability unaffected by the rapid changes of youth and the often exag-
gerated individual peculiarities of old age. As the specific differences
in the skull appear fully developed and most easily appreciated in
skulls between 40 and 70 mm. in basicranial length, I have selected
such specimens as norms to which other sizes may be more or less suc-
cessfully referred, pointing out the deviations, as shown in the available
material, under the headings of the various species. As the sexual
differences are comparatively slight they have been ignored in the key,
though it may be pointed out here that the adult males differ visibly
from the females in the tail extending considerably beyond the cara-
pace while in the females it about reaches the edge of the disk; in
possessing a smaller head and a greater expansion of the plastral cal-
losities.

It is finally to be emphasized that specimens may be met with
which depart so far from the norm that they defy positive identifica-
tion by the registered characters alone or in combination. Such cases
are not particularly rare among the Testudinata.

Key to young adult specimens of North American
soft-shelled turtles

a1 Neurals normally 8, separating all the pleurals; entoplastron
bent at an angle of about 100° or more; a central callosity nor-
mally on entoplastron; temporal fossa of skull small, the longest
diameter less than two thirds the long diameter of the orbit;
distance from orbit to tympanic cavity about equals diameter of
orbit; intermaxillaries (premaxillaries) narrowly touch or are
separated by the maxillaries. Nostrils rounded, septum between
them rather wide and without lateral projecting ridge; no tuber-
cles on the leathery disk of carapace or its anterior edge {mulica
group; subgenus Euamyda).

Amyda mutica, (p. 14)

a2 Neurals normally 7 (occasionally 8), last pair of pleurals in con-
tact; entoplastron bent at an angle of about 90° or less; no cal-
losity on entoplastron; temporal fossa on skull about equal to
or longer than long diameter or orbit; distance from orbit to

stejneger: American soft-shell turtles 13

tympanic cavity much greater than diameter of orbit; maxil-
laries in contact above intermaxillaries (premaxillaries). Nostrils
crescent-shaped, internal ridge projecting on each side from the
narrow septum between them; leathery disk of carapace with
tubercles at least on anterior edge.

b1 Length of inner bony choanae greater than their distance
from intermaxillary foramen; mandibular symphysis less or
equal to length of choanae {ferox group).
c1 wSculpture of bony carapace coarsely grained, usually with
numerous irregular, more or less continuous and anasto-
mosing longitudinal ridges. (Leathery carapace of young
with longitudinal rows of tubercles; coloration peculiar.

Amyda ferox, (p. 25)

c2 Sculpture of bony carapace finely grained ; (leathery disk
of carapace of young smooth ; coloration of upper surfaces
gray, more or less marked with small dusky ocellae, or
solid spots or lines; plastron uniform white).
d1 Tubercles on anterior edge of leathery carapace well
developed, normally triangular and pointed; distinct
tubercles covering anterior and posterior flaps.
e1 Bony carapace without raised bony knobs or
"warts"; (young with one marginal dusky line on
posterior part of leathery carapace).

Amyda spinifera, (p. 43)

e2 Adult with strong tubercles on the hind part of
carapace "supported there by prominent bony
warts of the bony plates" ; (in young margin of the
leathery carapace marked posteriorly by at least
two parallel dusky lines).

Amyda spinifera aspera, (p. 56)

d2 Tubercles on anterior edge of leathery carapace poorly

developed, short and bluntly rounded; tubercles on

both flaps, if present, quite minute, except in very old

specimens.

Amyda emoryi, (p. 65)

b2 Length of inner bony choanae equals their distance from in-
termaxillary foramen; mandibular symphysis longer than
length of choanae (agassizii group).

Amyda agassizii, (p. 72)

14 bulletin: museum of comparative zoology

The mutica group

The only species so far known in this group deviates in its characters
and their combination quantitatively more from any of the others in
the genus, so much in fact that it has been regarded by outstanding
zoologists such as Agassiz, Cope, True, and Baur as representing a
"good" genus. However, the differences are of a character that rather
suggest relationship with the ferox group than a separate phylogeny.
The difference in the number and relations of neurals does not seem to
have any genetic significance in the genus Amyda. The long drawn out
and slender snout with the occasional separation of the maxillaries at
the apex and the shortening of the temporal-tympanic region indicate
modifications due to some food specialization with which the absence
of the dermal ridge in the nostrils at the tip of the proboscis and the
thickening of the septum may be correlated.1 In fact, there is indica-
tion of the ridge inside, though not reaching the opening of the nostrils.
The extreme development of the plastral callosities is purely quan-
titative and is closely approached in old specimens of Amyda emoryi,
and a small callosity on the entoplastron is often observable in other
specimens of the genus. The more circular shape of the body outline
is rather a juvenile character with which the obtuse angle of the
epiplastral bones is correlated. In none of the characters does the A.
mutica show any approach to any other group in the genus, particu-
larly not to any of the Old World soft -shell turtles, so that there seems
to be no convenience in recognizing it as a separate genus.2

Amyda mutica3 (Lesueur)
Plates 2, 3, 4

1827. Trionyx muticus LESUEUR, Mem. Mus. Hist. Nat. Paris, 16, Dec.
1827, p. 263, pi. 7 (type-locality, Wabash River, New Harmony,
Indiana; type Paris Mus. No. 787; Lesueur, collector ).— LE CONTE,
Ann. Lye. Nat. Hist. New York, 3, 1830, p. 95.— GRAY, Syn. Rept.,

1 Holbrook, curiously enough, who especially called attention to the "two characters which
always exist", viz. the "total absence of spines or tubercles" and the "great difference of the
nostrils" in describing the latter speaks of them as "closely approximated" in both ferox (in
which he included spinifera) and mutica (North American Herpetology, Edit. 2, 2, p. 13 and
p. 19) while in reality thev "are widely apart" in the latter, as pointed out bv Agassiz (Contrib.
U. S. Nat. Hist., 1, p. 398).

'Should the statement made by Dr. Stockwell (Journ. Comp. Med. Surg., 9, 1888, p. 29)
be corroborated, viz., that "the marginal ossicles in A. mutica are rudimentary; in A. spinifer
altogether wanting," the question of the generic status of A. mutica might well be reopened.

3 Latin, muticus, docked, dehorned, with reference to the absence of spines on the anterior
edge of the leathery carapace.

stejxeger: American soft-shell turtles 15

pt. 1, 1831, p. 46; Cat. Tort. Brit. Mus., 1844, p. 50; Cat. Shields
Rept. Brit. Mus., p. 1, March 8, 1856, p. 69.— DUMERIL AND
BIBROX, Erp. Gen., 2, 1835, p. 482 (lapsus for Gymnopus).— HAR-
LAN, Med. Phys. Res., 1835, p. 159 (Ohio River and tributaries).—
WIED, Reise Nord-Amerika, 1, pt. 3, 1838, p. 140 (Pittsburgh, Pa.).
-HOLBROOK, North Amer. Herpet., 1 ed., 4, 1840, p. 17, pi. 2
(Mississippi and tributaries); 2 ed., 2, 1842, p. 19, pi. 2 (Mississippi
and tributary streams).— DE KAY, Zool. New York, Rept., 1842,
p. 7 (Ohio River).— TROOST, Seventh Geol. Rep. Tennessee, 1844,
p. 39 (Tennessee; milieus misprint). — DUMERIL, Cat. Meth.
Rept. Mus. Paris, pt. 1, 1851, p. 22 (types).— STRAUCH, Mem.
Acad. Sci. St, Petersbourg, ser. 7, 5, No. 7, 1862, p. 174; 8, No. 13,
1865, p. 125; 38, No. 2, 1890, p. 118.— HOY, Geology of Wisconsin,
1, 1883, (p. 423) (Wisconsin).— BOULENGER, Cat. Chel. Brit.
Mus., 1889, p. 260, fig. 68 (Mississippi, Ohio and Saint Lawrence). —
HAY, Indiana Geol. 17 Rep., 1892, p. 551; Batr. Rept. Indiana,
1893, p. 143 (Indiana: Delphi; Madison; Terre Haute. Illinois:
Mt. Carmel).— HURTER, Trans. Acad. Sci. St. Louis, 6, 1892, p.
259 (Missouri: Mississippi River near St. Louis).— SIEBENROCK,
Sitz. Ber. Akad. Wiss. Wien, Math.-Nat. KL, 111, 1902, p. 822, fig. 5
(plastron); Zool. Jahrb. Suppl., 10, pt. 3, 1909, p. 605; Ann. Naturh.
Hofmus. Wien, 27, 1913, p. 214, sep. p. 44 (plastron); Verh. Zool.
Bot. Ges. Wien, 73, Aug. 1923, p. 192.— DITMARS, Reptile Book,
1907, p. 78, pi. 27, low. fig. (St. Louis, Mo.).— OVER, South Dakota
Geol. Nat. Hist. Surv. Bull. 12, Oct. 1923, p. 18, pi. 7 (Missouri Riv.
and eastward, South Dakota).

Gymnopodus muticus DUMERIL, Arch. Mus. Hist. Nat. Paris, 7,
1855, p. 203.

Gymnopus muticus DUMERIL and BIBRON, Erpet, Gen. 9, 1854,
p. 236.— WIED, Nova Acta Leopold.-Carol., 32, pt. 1, 1865, p. 54
(Ft. Mackenzie, Missouri River, 6-8 miles below Cedar Isl., South
Dakota).

Amyda mutica AGASSIZ, Contr. Nat. Hist. United States, 1, 1857, p.
399; vol. 2, pi. 6, figs. 6-7 (Lake Erie and Ontario; Delphi, Ind.;
Burlington, Iowa; Osage River, Missouri; Alleghany Riv., Pa.). —
GRAY, Suppl. Cat. Shield Rept. Brit. Mus., p. 1, 1870, p. 95.—
COPE, Bull. U. S. Nat. Mus., No. 1, 1875, p. 41 (middle and northern
tributaries of Mississippi and the St. Lawrence Riv.). — JORDAN,
Man. Vert. North. United States, ed. 3, 1880, (p. 168); ed. 8, 1899,
p. 206 (Canada to Ohio River, and N. W.).— CRAGIN, Trans. Kan-
sas Acad. Sci., 7, 1881, p. 116 (Kansas: Manhattan; Blue and Kansas
Rivers).— SMITH, Rep. Geol. Surv. Ohio, 4, 1882, p. 668 (Ohio
River, Ohio).— TRUE, Bull. U. S. Nat. Mus., No. 24, 1883, p. 28
(Madison, Ind.; Mt. Carmel, 111.; St. Louis, Mo.; Ft. Smith, Ark.);
Fish Industr. United States, sect. 1, 1884, p. 152.— HOY, Geol.

16 bulletin: museum of comparative zoology

Wisconsin Survey 1873-1879, 1, 1883, p. 423 (tributaries to the Mis-
sissippi in Wisconsin). — DAVIS and RICE, Illinois State Lab. Nat.
Hist. Bull. No. 5, 1883, p. 52 (North of Ohio River); Bull. Chicago
Acad. Sci., 1, No. 3, 1883, p. 32 (Illinois).— BAUR, Zool. Anz., 10,
1887, p. 99 (descr. plastron) ; Amer. Natural., 22, 1888 (1122); Proc.
Amer. Philos. Soc, 31, 1893, (p. 220).— STOCKWELL, Journ.
Comp. Med. Surg., 9, No. 1, Jan. 1888, p. 29.— HIGBY, Trans.
Wisconsin Acad. Sci., 7, 1889, p. 159 (Wisconsin: western half of
state).— GARM AN, H., Bull. Illinois Lab. Nat. Hist., 3, 1892, p.
247 (throughout Illinois: Mackinaw Creek, Woodford Co.; Quincy;
Illinois Riv., Peoria; Wabash Riv., Mt. Carmel; Ohio Riv., Cairo). —
RHOADS, Proc. Acad. Nat. Sci. Philadelphia, 1895, p. 404 (Ten-
nessee).— GAGE, Trans. American Microsc. Soc, 17, 1895, (p. 185)
(embryology).— SMITH, Proc. Linn. Soc. New York, 1898-1899,
No. 11 (1899) p. 24 (not near New York City).^-McLAIN, Notes
Coll. Rept. Arkansas, 1899, p. 1 (Ft. Smith, Ark.).— ATKINSON,
Ann. Carnegie Mus. Pittsburgh, 1, 1901, p. 154 (Allegheny Co.,
Pa.).— PAULMIER, Bull. 51, New York State Mus., 1902, p. 392
(probably in northern New York State). — MORSE, Proc. Ohio
Acad. Sci., 4, pt. 3, Spec. Pap. No. 9, 1904, p. 138 ("not recorded for
Ohio")- — ?NASH, check list of Vertebrates of Ontario, Batrachians,
Reptiles, Mammals, 1906, p. 17 (Ontario, Lake Erie, rare).
SURFACE, Zool. Bull. Pennsylvania Dep. Agric, 6, nos. 4-5,
Sept. 1, 1908, p. 119 (Pennsylvania, liable to be found in Erie
County and tributaries of the Ohio). HURTER and STRECKER,
Trans. Acad. Sci. St. Louis, 18, no. 2, 1909, p. 21 (Arkansas:
Ft. Smith; Pink Bluff; Little Rock). — HURTER, Herpet. Mis-
souri, 1911, p. 249 (Mississippi, Osage, Gasconade and Meranac
Rivers).— CLARK and SOUTHALL, Rep. U. S. Comm. Fish.,
1919 (1920) App. No. 7, p. 15 (economics). — MULLER,
Amer. Midland Natural., 7, no. 6, Nov. 1921, p. 180 (Iowa:
Fairport; habits). — BLANCH ARD, Occas. Pap. Zool. Mus.
Univ. Michigan, No. 117, July 6, 1922, p. 18 (Trotter's Landing,
Benton Co., Tenn.).— WEED, Copeia, No. 116, March 15, 1923, p.
48 (Meredosia, Ills.).— PRATT, Man. Vert. United States, 1923, p.
249.— STRECKER, Baylor Univ. Bull., 27, No. 3, Sept. 1924, p. 47
(Little Rock, Ark.); Contr. Baylor Univ. Mus., No. 7, July 15, 1926,
p. 7 (Neches River, Henderson Co., Texas, probably). — MYERS,
Proc. Indiana Acad. Sci., 35, 1926, p. 292 (Indiana).— STRECKER
and FRIERSON, Contr. Baylor Univ. Mus., No. 5, May 15, 1926,
p. 10 (Caddo and De Soto Parishes, La.).— ORTENBURGER,
Proc. Oklahoma Acad. Sci.,' 6, pt. 1, 1926, p. 100 (McCurtain and
Pushmataha Cos., Oklahoma).— STRECKER and WILLIAMS,
Contr. Baylor Univ. Mus., No. 12, Dec. 27, 1927, p. 16 (Christoval,
Tom Green Co., Texas); No. 17, Oct. 20, 1928, p. 19 (Bowie Co.: "no

stejneger: American soft-shell turtles 17

doubt")-— BURT, Occas. Pap. Mus. Zool. Univ. Michigan, No. 189,
Dec. 12, 1927, p. 9 (Manhattan, Riley Co., Kansas) ; Amer. Midland
Natural., 16, No. 3, 1935, p. 321 (Barber and Reno Cos., Kansas).—
POPE and DICKINSON, Bull. Publ. Mus. Milwaukee, 8, No. 1,
April 3, 1928, p. 82, pi. 21, figs. 5-6 (Wisconsin: Mississippi River
counties; Crawford and Pepin Cos.). — JORDAN, Man. Vert.
Northeast, U. S. (13 ed.) 1929, p. 254 (Middle and Northern tribu-
taries, Miss, and St. Lawrence rivers.) — ORTENBURGER and
FREEMAN, Publ. Univ. Oklahoma Biol. Surv., 2, 1930, p. 188
(Oklahoma: Alfalfa and Comanche Cos.). — WALKER, Copeia,
1931, No. 1, p. 12 (Scioto and Brown Cos., Ohio).— BOYER and
HEINZE, Trans. Acad. Sci. St. Louis, 26, No. 4, Apr. 1, 1934, p. 199
(Missouri: Jefferson Co.: Meranac River). — RUST, Blatt. Aquar.
Terrarienk., 45, 1934, p. ; sep. p. 12.— TAYLOR, Univ. Kansas
Sci. Bull., 22, No. 11, Apr. 15, 1935, p. 218 (Arkansas: Duvall Bluff,
Prairie Co.; Lewisville, Lafayette Co.).— DELLINGER and
BLACK, Occas. Pap. Univ. Arkansas, No. 1, June 1938, p. 46 (Ar-
kansas: Garland, Jefferson, Lafayette, Prairie, Pulaski, Sebastian,
Franklin and Lawrence Cos.). — SOLA, Bull. New York Zool. Soc,
34, No. 5, Sept.-Oct., 1931, pp. 134, 142, 155, fig. 7 upper (western)
part of Pennsylvania along Lake Erie). — CAHN, Illinois Biol.
Monogr., 16, Nos. 1-2, Aug. 31, 1937, p. 176; pis. 24, 30 fig. a; map
19(Illinois).— CONANT, Amer. Midland Natural., 20, No. 1, July,
1938, p. 154, pi. 21, fig. 1 (left), fig. 2 (Ohio: Scioto, Muskingum and
Ohio Rivers; map).— PARKER, Rep. Reelfoot Lake Biol. Sta., 3,
Jan. 1939, p. 88 (Tennessee: Reelfoot Lake, nowhere abundant). —
WELTER and CARR, Copeia, 1939, No. 3, Sept. 9, p. 130 (Ken-
tucky, east.: Triplet Co.; Fox, Fleming Co.; rare). — LOGIER, Roy.
Ontario Mus. Zool. Hanb. No. 4, p. 57 (Ontario: Lake Erie, probably
misidentified).— GENTRY, Rep. Reelfoot Lake Biol. Sta., 5, Jan.
1941, p. 75 (Tennessee: Pickett Co.); Journ. Tennessee Acad. Sci.,
16, No. 3, 1941, p. 332 (Tennessee: Obey River, Pickett Co.).—
ANDERSON, Bull. Chicago Acad. Sci., 6, No. 11, 1942, p. 219
(Missouri: Jackson Co.: Fry's Lake).— PETERS, Copeia, 1942, No.
3, Oct. 8, p. 183 (Illinois: Cumberland Co.).
1864. Potamochdys ? microcephala GRAY, Proc. Zool. Soc. London, 1864
(p. 87) (type-locality, Sarawak, Borneo!!; type in British Museum).
Callinia microcephala GRAY, Proc. Zool. Soc. London, 1869 (p.
222); 1873, p. 62, fig. 11; Suppl. Cat. Shield Rept. Brit. Mus., pt. 1,
1870, p. 108; Hand-list Shield Rept. Brit. Mus., 1873, p. 83.

Types. In the Musee d'Histoire Naturelle at Paris, No. 787 is
a shell on the plastron of which is written: "Trionix mutica Lesueur.

Wabash River par moi aout 1S27", and on a paper label pasted

on the underside of the stand: "Tortue qui n'a point le bord desou

18 bulletin: museum of comparative zoology

disque spineaux & que j'ai designee sou le nom de mutica dans

precedente note accompagniee de figure que je vous ai fait passer |?]
C. A. Lesueur." It is designated as the type on the printed label. The
bony carapace is 106 mm. in length and 107 mm. in width. A pair of
fontanelles between first pair of pleurals ; 8 neurals separate the pleurals.

No. 788 is a cotype retained by Lesueur until 1844 when it was ac-
quired by the Museum. It is a larger, mounted specimen, the bony
carapace measuring 134 mm. in length, 147 mm. in width; fontanelles
obliterated; 8 neurals, but last pair of pleurals broadly in contact.
Underside of stand inscribed "Gymnopus muticus Lesueur. Wabash.
Acquis de Mr. Lesueur 1844" (Lesueur died December 12, 1846).

The so-called "Spineless" or "Brown Softshell Turtle", the smaller
of our American species, may be easily recognized by the characters
given in the key. In the adolescent and adult carapaces the absolute
absence of tubercles on the anterior edge of the leathery disk will
identify the A. mutica even when the head is missing or the proboscis
is so mutilated as to defy examination. At the stage when the very
young specimens of all the species are nearly circular in outline and the
tubercles on the anterior edge of carapace of those species normally
possessing them may in some individuals be so indistinct as to be
doubtful, the oval nostrils without the septal tubercle will positively
identify A. mutica.

The other differential characters keyed are not always to be relied
on because of individual variation. Thus while normally the number of
pleurals is 7 pairs, separated the entire length of the bony carapace by
a series of 8 neurals, there are many and significant exceptions. Thus,
as already mentioned, the mounted cotype of the species in the Paris
Museum has the last pair of pleurals broadly in contact. Similarly the
U.S.N.M. 102910 has the eighth pair of pleurals in contact for at least
half their length; No. 54734 has all the pleurals separated by the 8
neurals, but it has only 6 pleurals on the right side, against 7 on the
left; U.S.N.M. 92605, 95134 and 029261 have only 7 neurals and 7
pairs of pleurals and the last pair is broadly in contact behind the
neurals. While visiting the Museum of Comparative Zoology during
the early days of these investigations, I examined two specimens, both
unfortunately at that time without numbers and locality, one adoles-
cent with the seventh pair of pleurals in contact behind the neurals,
while the second, an adult skeleton with a bony carapace measuring
140 mm. in length and 150 mm. in width, was still more abnormal
having 8 pleurals on the right side, against 7 on the left, and with 9
neurals separating all the pleurals.

stejneger: American soft-shell turtles 19

The greater angular width of the entoplastron is quite characteristic
of this species, but it is sometimes questionable in application because
of difficulty of exact measurement. However, it is useful where other
characters are irregular or in case one has to identify a disassociated
plastron or a single bone. The great extension of the plastral callosities
in the males is also a character of value, though the callosities on old
male A. emoryi may reach similar proportions on the hyo-, hypo-, and
xiphi-plastra. A central callosity is normally present in A. mutica on
the entoplastron though exceptional in the other species. Small callosi-
ties on the epiplastra are not uncommon in A. midica, though rare in
the others.

The sutural meeting of the maxillaries on the upper side of the
snout above the premaxillary (intermaxillary) is one of the characters
of the trionychid skulls, and is probably a normal or at least original
condition in A. mutica, but as noted above the extreme tapering and
hence weakening of the snout in this species results in the frequent
loss of these parts in the preparation of the skulls. In my series of
A. mutica skulls there are only four perfect specimens and in one of
these, No. 102677, the maxillaries are plainly in contact on the upper
side, above the premaxillary, while in the others, Nos. 53521, 54733 and
029261, the maxillaries are separated by the premaxillary.

A negative character of the young A. mutica is the absence of
ocellated or solid black rounded spots on the carapace, and of a defined
angular figure on top of the snout at the base of the proboscis. In a gen-
eral way the coloration is less distinctive than in the other species. The
yellowish margin of the leathery carapace seems to be definitely wider.

The coloration and pattern in this as well as the other species of
the genus becomes gradually more obscure with age, and varies in-
dividually as well as locally according to environmental conditions.
It may therefore not be unwelcome if I include a few color descriptions
of living or freshly killed specimens which have come under my ob-
servation when comparison with Ridgway's "Nomenclature of Colors
for Naturalists" (1886) was possible.

On September 4, 1934, the National Museum received from C. R.
Rogers two live specimens taken in Medicine Lodge River, one mile
SW of Lake City, Barber Co., Kansas. Description was at once made:
U.S.N.M. No. 95185, young adult female (leathery disk about 200
mm.). Iris bright "buff", the ring nicked slightly in front and behind
by a small blackish spot. — General color above nearly uniform
"tawny" with very faint mottlings of lighter "tawny-ochraceous" and
darker "raw umber", especially on the posterior flap, on which a faint

20 bulletin: museum of comparative zoology

submarginal dusky line borders the pale margin which is lightly
suffused with "rufous"; top and sides of head like back with a sharply
defined band of "ochraceous buff" narrowly edged with dusky; the
band continues — though fainter — anteriorly through the eye on to
the canthus rostralis converging towards the base of the proboscis
without meeting that of the other side; from the side of the occiput
indication of the band — though more irregular — on the side of the
neck; underside with a fine network of red blood vessels shining
through imparting a pinkish tinge to the soft parts which fades
gradually through "lavender" and "pearl blue" into "china-blue" on
the palms, soles and digits merging on the underside of the webs into
"tawny ochraceous" exteriorly and more pinkish interiorly; throat and
chin like the soles; lower lips whitish; callosities pale "fawn-color"
tinged centrally with blueish.

The other specimen, No. 95186, is a much smaller male (leathery
carapace length about 135 mm.) (pi. 3). Iris, a pale yellowish ring, but
the black spots are somewhat larger than in the older specimen.
Colors are also essentially like the latter, but the "ochraceous buff"
postocular band has the edges even better defined and on the side of
the neck joins the pale color of the underside which is sharply set off
from that of the upper side and extends onto the upper lip; the band
is only indicated in front of the eye by a small elongated triangular
spot of pale "ochraceous".

About the same time the National Museum received two live speci-
mens from J. H. Hall, collected in Mississippi (Marion County, Colum-
bia), both females. No. 95133, the larger one (disk approximately 185
mm. long). Iris bright buff yellow with a black horizontal bar. General
color above "clay color" with irregular blotches of pale "raw sienna";
marginal dark ring on carapace broken, faint, "hair-brown"; upper
side of neck washed with "tawny ochraceous"; postocular stripe dull
"buff -yellow", edged with dusky ("hair-brown"); front legs above as
well as dorsal and lateral surface of neck sprinkled with small dusky
spots; hind feet pale "olive yellow" with slightly larger and darker
dots and marblings; webs verging on "clay color"; underside of plas-
tron "flesh color", which on the white ground of the legs changes into
"pale blue" and on the soles and front feet verges on "heliotrope
purple"; underside of webs pinkish towards the edge; claws horny
white; callosities (none on epiplastra) "vinaceous-cinnamon". No fork
figure on top of snout; no spots or definite dusky markings on soles.

The smaller specimen, No. 95134 (leathery disk approximately 155
mm.) essentially as the larger one.

stejneger: American soft-shell turtles

Table 2
Cranial measurements of mutica in millimeters

CD
M

of
PE

I— 1

l-c

e*-

a
'3

o
1— 1

o
"a;

03
CO

53
«

CO
0)

a
a

CD
bfl

<M
iO

41.0
11.0

i— i

CO
CM
OS

42.0
12

i— i

43.0

48.0
14.5

OS
(N

T— t

Basicranial length

40.0
11.5

42.8

Tip of snout to orbit

12.3

Orbit, horizontal diameter

8.0

8.5

9.0

9.5

7.0

8.6

Orbit to tympanic cavity

8.0

8.5

9.0

10.0

9.0

8.8

Temporal fossa, longest di-

ameter

5.0

4.-r

6.0

5.0

6.0

5.0

5.3

Interorbital width

2.0

..0

2.0

3.0

2.0

2.1

Maxillary alveolar surface,

width

2.0

2.5

2.0

1.5

2.0

2.5

2.0

2.1

Internal choanae, length

6.0

7.0

6.5

6.0

6.5

7.0

6.7

Choanae to intermaxillary

foramen

4.0

4.5

5.0

4.0

4.2

Intermaxillary foramen,

length

6.0

6.5

8.0

6.5

6.7

Mandibular symphysis,

length

8.0

8.5

8.0

9.0

10.0

8.0

8.6

Mandibular alveolar sur-

face, width

The colors of a freshly killed male specimen (leathery disk 204 mm.
long) collected by Dr. C. E. Burt in Kansas, Reno County, 6 miles E.
of Turon (U.S.N.M. No. 95259) were as follows: Ground color "broc-
coli brown" mottled with numerous very irregular and more or less
anastomizing, ragged-edged "sepia" spots occupying as much space as

22 bulletin: museum of comparative zoology

the ground color; posterior edge of disk pale tinged with "burnt um-
ber"; no submarginal blackish line; upper soft parts same broccoli
brown with very small and faint irregular "sepia" dots; a very pale
"russet" band, raggedly edged with blackish, from canthus rostralis
through eye and over ear slanting on side of neck halfway down the
neck ; pale lines in front of eyes do not meet those on snout nor is there
trace of a connecting line forming fork or triangle at base of proboscis ;
underside whitish with a faint glaucous blue gray tinge on throat,
underside of neck and feet; no dusky markings on feet; callosities, in-
cluding the small central one on entoplastron, pale blueish "plumb-
eous".

Geographical Distribution

Mississippi River and tributaries; north to South Dakota and
Minnesota; east to western Pennsylvania; west to Kansas, Oklahoma
& Texas.

Recorded from northern localities on the Trinity, Brazos and
Colorado Rivers.

Agassiz (Contr. Nat. Hist. U. S., 1, p. 404, footnote) writes, "De-
Kay's Trionyx ocellatus is Amyda mutica," a statement which seems
to be erroneous. The reference to DeKay's Trionyx ocellatus appears to
be the following note in his Zoology of New York (pt. 3, 1842, p. 7)
under Trionyx ferox [DeKay= T. spiniferus]: "The description given
above [p. 6: "anterior margin in the adult with numerous pointed
tubercles, which may be faintly and distantly traced in the young"]
was taken several years since, from a specimen obtained in the Mohawk
River. . . . The specimen, as I then thought, varied so much from any
description of the ferox within my reach, that I considered it to be new,
and named it ocellatus." The description of the Mohawk River speci-
men is clearly that of an adult A. spin if era, and cannot be taken as the
record of A. mutica in the Mohawk River. DeKay's mention that he
suspected ferox and muticus to be identical probably caused Agassiz's
statement.

List of specimens in U. S. National Museum

4783 (2) <? a<

id., d" adol. ?

7646 d" adol.

Mo., St. Louis

Engelmann

7647 juv.

U tl

7655 juv.

Miss., Monticello

Helen Tennison

7659-60 (2)

111.

Kennicott

7746

Ark., Arkansas R. near

Ft. Smith

Lt. Whipple

stejneger: American soft-shell turtles

8337 9

ad.

Indiana, Madison

9615 c?

adol.

111., Mt. Carmel

Mrs. M. E. Turner

9646 d1

adol.

11 tt

Robert Ridgway

May

1878

9647 d" adol.

II tt

(«

9650 <? adol.

it a

9651 rf" adol.

t; it

9727 9

ad.

it ll

J. Schneck

11629 juv.

11630 9

adol.

12794 (2.

I juv.

111., Mt. Carmel

J. Schneck

13549

14780 juv.

Mo., St. Louis

J. Hurter

Dec.

20, 1887

19626 9

ad.

19627 9

adol.

21418 9

22629 juv.

Tex., Sabine R., 5 mi. S.

Jordan & Gilbert

1884

Longview.

029261 ad

Dr. G. Baur

45735-8

Iowa, Fairport, Missis-

sippi R.

Snyder

June

2, 1916

52528 <? ad.

Kansas

Roy L. Moodie

52116-8

juv.

111., Olney

Robert Ridgway

53521 9

ad.

Iowa, Fairport

Bur. Fisheries

Apr.

24, 1911

54733 9

ad.

ii it

J. Snyder

54734 9

ad.

it tt

Aug.

8, 1916

54742

" (Missis-

sippi R.)

May

8, 1916

55525

Mo., St. Louis

J. Hurter

Mar. 23, 1907

55526

May

14, 1913

55527

Ark., Jeff. Co.

1899

55528

" Little Rock

June

1, 1900

55600

Tex., McLennan Co.

J. Hurter

1896

59267 9

ad.

Mo., Alexandria

E. Stringham

June

5, 1916

59268-9

Minn., Homer

F. Schrader

Sept.

8, 1916

59276

111., betw. Warsaw &

Hamilton

E. Stringham

Aug.

23, 1916

59278 9

ad.

Mo., Alexandria

June

5, 1916

59281

Iowa, Keokuk

Aug.

16, 1916

59282

tt it

June

7, 1916

59283

it it

July

1, 1916

59284

it it

June

7, 1916

59982

"Central U.S."

O. P. Hay

60054-6

Iowa, Fairport

Bur. Fish.

60561

Mo. St. Louis

J. Hurter

bulletin: museum of comparative zoology

71547

Okla., 6 mi. E. Ingersoll

A. I. Ortenburger

92605 cf adol.

Miss., Greenville

S. F. Hildebrand

May 29, 1933

95133-4 9 adol. Miss., Columbia

J. H. Hall

Aug. 1934

95185 9 ad.

Kans., 1 mi. S.W. Lake

City

C. R. Rogers

Aug. 31, 1934

95186 d1 juv.

Kans., 1 mi. S.W. Lake

City

95259 a*

Kans., 6 mi. E. Turon

C. E. Burt

May 25, 1934

95260 9 adol.

U U {{

95415 9 adol.

" 3 mi. S.E. Oxford

1934

100422 9 adol.

La., Rayville

Aug., 1935

100813 pull.

Kans., Wakeeney

O. P. Hay

102612-3

111., 5 mi. S. Savannah

P. Bartsch

July 29, 1907

102677 9 ad.

Tenn., Reelfoot Lake

W. M. Perrygo &

C. Lingebach

May 6, 1937

102910 9 ad.

it it a

W. M. Perrygo &

C. Lingebach

May 8, 1937

115939

[Miss.]

[B. C. Wailes]

The ferox Group

Although quite distinct, the three species included form a rather
close group chiefly characterized by the uniformity of their skulls.
Common for all three is the large opening of the inner choanae with the
concomitant shortness of the distance of the latter from the inter-
maxillary foramen, which distinguish them from the agassizii group,
while the normal proportion of the preorbital region sets them off from
the mutica group. This statement may seem strange in view of the
opinion of many early competent herpetologists who refer to the species
of the ferox group under two different generic terms, a situation caused
by the confusion of the identity of the specimens which served as basis
for the generic concept, as will be shown later on.

Externally the ferox group differs from the mutica group by the
crescentic shape of the nasal openings, a character shared by all the
other species of the genus, and by the presence of the dermal tubercles
on the carapace. From the agassizii group, however, there is no obvious
external character by which to distinguish them as a group.

Within the group, the species from which its name is taken (because
the oldest one known) is most easily identified in the adult stage from
the other American species by the coarseness of the sculpture of its
bony callosities (pis. 9, 10), and in the early juvenile stage by its
unique coloration (pi. 19). The greater extension of the plastral
flap anteriorly beyond the carapace is also a noteworthy feature.

STEJXEGER: AMERICAN SOFT-SHELL TURTLES 25

Amyda ferox1 (Schneider)

Plates 5-10

1783.— Tesludo ferox SCHNEIDER, Naturg. Schildkr., p. 330 (Savannah
River, Georgia; type in Brit. Mus.; Dr. A. Garden, collector) (based
on Pennant, Philos. Trans., 61, pt. 1, p. 266).— GMELIN, Syst.
Nat., 1, pt, 3, 1789, p. 1039.— SCHOEPFF, Naturg. Schildkr., pt.
5, 1795, p. 102; Hist. Testud., pt. 5, 1795, p. 88 (based on Pennant).-
SHAW, Gen. Zool., 3, 1802, p. 64, pi. 17.— LATREILLE, Hist. Nat.
Rept,, 1, 1801, p. 165 (based on Pennant).— DAUDIN, Hist. Nat.
Rept., 2, 1802, p. 69 (based on Pennant).
Trionyx ferox SCHWEIGGER, Konigsberg. Arch. Naturw. Math., 1,
1812, pt. 3, p. 285 (Carolina and Florida) ; pt. 4, p. 363; Prodr. Mon.
Chelon., pt. 1, 1814, p. 15.— MERREM, Tent. Syst. Amph., 1820,
p. 20. — SAY, Journ. Acad. Nat. Sci. Philadelphia, ser. 1, 4, pt. 2,
1825, p. 218 (part: Carolina; Georgia). — HARLAN, Journ. Acad.
Nat. Sci. Philadelphia, 6, Feb. 1827, p. 32 (part: many of the rivers
of the southern states, not observed to exist further south [north ?]
than South Carolina) ; Medic. Phys. Res., 1835, p. 158.— Le CONTE,
Ann. Lye. Nat. Hist., New York, 3, 1830, p. 93 (part: Rivers of
Georgia and Florida, north to Savannah). — GRAY, Synops, Rept.,
1831, p. 45 (part); Cat. Tort. Brit. Mus., 1844, p. 49 (part); Cat.
Shield Rept. Brit. Mus., pt. 1, March 1856, p. 68 (part); HOL-
BROOK, North Amer. Herpet., ed. 1, 4, 1840, p. 9, pi. 1 (part:
Savannah and rivers emptying into northern border of Gulf of
Mexico); ed. 2, 2, 1842, p. 11, pi. 1 (part); in White's Statistics of
Georgia, 1849, Fauna and Flora, p. 13 (Georgia).— STRAUCH,
Mem. Acad. Sci. St. Petersbourg, ser. 7, 5, no. 7, 1862, p. 173
(part); 8, no. 15, 1865 p. 122. — BOULENGER, Cat. Chel. Brit.
Mus., 1889, p. 259 (Georgia, Louisiana).— DITMARS, Rept. Book,
1907, p. 74, pi. 26, lower fig. (Georgia, Florida, Louisiana). — SIE-
BENROCK, Sitzungsbr. Akad. Wiss. Wien, Math. Nat. CI., 91, pt.
1, Oct. 1902, p. 829; Zool. Jahrb. Suppl., 10, pt. 3, 1909, p. 603
(Georgia, Florida west to Louisiana); Verh. Zool. Bot. Ges. Wien,
73, Aug. 1923, p. 181. — BRIMLEY, Proc. Biol. Soc. Washington,
23, 1910, p. 18 (Georgia: MimsviUe; Florida: Orlando; Belleaire;
Green Cove Springs; St. Petersburg).— DECKERT, Copeia, No.
54, Feb. 17, 1918, p. 31 (Jacksonville, Duval Co., Fla.).— FLOWER,
Proc. Zool. Soc. London, ser. A, 1937, pt. 1, Apr. 15, pp. 16, 37
(age: 25 years + ). — POPE, Turtles of the United States and
Canada, 1939, p. 303, pi. figs. 94-97 (Southeastern Atlantic and
Gulf Coastal Plain).

1 Ferocious, the specific name evidently refers to the following sentence in Dr. Garden's
original description: "As the animal is very fierce, when it is attacked or disturbed, it often
raises itself on its legs, and will leap forward to bite its disturber or enemy, which it does with
great fury and violence".

26 bulletin: museum of comparative zoology

Amyda ferox OKEN, Lehrb. Zool., 2, 1816, p. 348.— STEJNEGER
and BARBOUR, Check List North Amer. Amph. Rept., ed. 1, 1917,
p. 124 (South Carolina to Florida and Louisiana); ed. 2, 1923, p. 140;
ed. 3, 1933, p. 153; ed. 4, 1939, p. 171.— SHUFELDT, Aquat. Life,
6, 1920, p. 27 (Georgia to Florida and Louisiana; habits). — LODING,
Alabama Mus. Nat. Hist., Paper No. 5, Sept. 1922, p. 47 (Fig
Island, Mobile Co., Ala.).— PRATT, Man. Vert. United States,
1923, p. 250.— WRIGHT, Ecology, 7, Jan. 1926, p. 84, pi. 5, fig. 3
(Okefinokee region, Ga.).— CORRINGTON, Copeia, 1927, No. 165,
Dec. 23, p. 101 (Pascagoula Swamp, near Biloxi, Harrison Co.,
Miss.).— PICKENS, Copeia, 1927, No. 165, Dec. 23, p. 43 (South
Carolina).— JORDAN, Man. Vert. Northeast U.S., (13 ed.) 1929, p.
255 S.C. to Fla. & La.— CONANT, Bull. Antiven. Inst. America, 4,
No. 3, Dec. 1930, p. 63 (Florida: Seminole; 15 m. E. of Sarasota; 18
m. S. of Ft. Myers).— HALTOM, Alabama Mus. Nat. Hist., Pap.
No. 11, 1931, p. 141 (pi. 39) (Alabama part, Mobile Co., Fig IsL). —
VAN HYNING, Copeia, 1933, No. 1, Apr. 5, p. 7 (Alachua Co.,
Fla., moderately common). — DeSOLA and ABRAMS, Copeia,
1933, No. 1, Apr. 3, p. 12 (Okefinokee Swamp, Ga.).— ALLEN,
Amer. Mus. Novit., No. 542, June 20, 1932, p. 20 (Mississippi: Han-
cock Co.).— RUST, Blatt. Aquar. Terrarienk., 45, 1934, sep. p. 12.

Platypeltis ferox FITZINGER, Syst. Rept., 1843, p. 30.— AGASSIZ,
Contr. Nat. Hist. United States, 1, 1857, p. 401 (southern states,
Georgia to western Louisiana). — GRAY, Proc. Zool. Soc. London,
1869 (p. 214); 1873 (p. 58) (part).— TRUE, Fish. Industr. United
States, sect. 1, 1884, p. 152.— BAUR, Proc. Amer. Philos. Soc, 31,
1893, p. 220— LONNBERG, Proc. U. S. Nat. Mus., 17, 1894, p.
317 (southern Florida).— WRIGHT and BISHOP, Proc. Acad. Nat.
Sci. Philadelphia, vol 67, 1915, p. 119, pi. 1, figs. 1, 2, 4; pi. 2, fig. 6
(Okefinokee Swamp, Georgia).

Aspidonectes ferox COPE, Bull. U. S. Nat. Mus., No. 1, 1875, p. 51
(Georgia to western Louisiana), — TRUE, in H. S. Thompson, South
Carolina, 1883, p. 238 (South Carolina); Bull. U. S. Nat. Mus., No.
24, 1883, p. 29 (part: Palatka, Putnam Co., Fla.; Charleston, S. C).
—DAVIS and RICE, Illinois State Lab. Nat. Hist. Bull. No. 5,
1883, p. 52 (Georgia to western Louisiana).— SHUFELT, Rep. U.S.
Nat. Mus. 1892 (1893) p. 32 (cast U.S.N.M. No. 8899).— BAUR,
Amer. Natural., 22, 1888 (p. 1121).— COKER, Bull. North Carolina
Geol. Surv., 14, 1906, p. 66 (North Carolina, introduced ?).— GOFF
and GOFF, Copeia, 1935, No. 3, Oct. 15, p. 156 (Griffin, Lake Co.,
Fla.; incubation).

1788.— Testudo mollis LACEPEDE, Hist. Nat. Quadr. Ovip. Serp., 1, Synops.
Meth. Quadr. Ovip., tab. between pp. 618 and 619 (based on Pen-
nant).— BONNATERRE, Tabl. Enc. Meth. Erpet., 1789, p. 25.

stejneger: American soft-shell turtles 27

1795.— Testudo (ferox ?) verrucosa SCHOEPFF, Naturg. Schildkr., pt. 5, p. 105
(based on Bartram's Trav. North and South Carolina, 1791, p. 177)
(type locality, eastern Florida); Hist. Testud., pt. 5, 1795, p. 90.

1801.— Testudo bartrami DAUDIN, Hist. Nat. Rept., 2, p. 74 (based on Bar-
tram, Voy. Amer. Septentr., vol. 1, p. 307).— HARPER, Amer. Mid-
land Natural., 23, No. 3, May 1940, p. 717 (Halfway Pond, Putnam
Co., Florida, "restricted type locality").
Trionyx bartrami GEOFFROY-ST. HILAIRE, Ann. Mus. Hist. Nat.
Paris, 14, Aug. 1809 (p. 18).— Le CONTE, Ann. Lye. Nat. Hist.
New York, 3, 1830, p. 96 (St. John's River, Fla.).

1809 1— Trionyx carinatus GEOFFROY-ST. HILAIRE, Nouv. Bull. Soc.
Philom. Paris, 1, No. 22, July 1809, p. 365 (type locality unknown;
type in Paris Mus.); Ann. Mus. Hist. Nat. Paris, 14, Aug. 1809, p.
14, pi. 4.

1809.— Trionyx georgianus GEOFFROY-ST. HILAIRE, Nouv. Bull. Soc.
Philomat. Paris, 1, No. 22, July 1809, p. 367 (substitute name for
T. ferox "Pennant").

1809.— Trionyx georgicus GEOFFROY-ST. HILAIRE, Ann. Mus. Hist. Nat-
Paris, 14, Aug. 1809, p. 17 (variant).

1812 ?.— Trionyx brongniarti SCHWEIGGER, Konigsberg. Arch. Naturw-
Math., 1, p. 288 (substitute name of Trionyx carinatus); Prodr. Mom
Chelon., 1814, p. 18.

1835. — Gymnopus spiniferus DUMERIL and BIBRON, Erpet. Gen., 2, p. 477
(part: Rivers of Georgia and Florida).

1835.— Trionyx harlani "Bell" HARLAN, Medic. Phys. Research, p. 159
(type locality, East Florida; type in "Mus. of Bell; Lond.").—
DeKAY, Zool. New York, Rept., 1842, p. 7 (East Florida).

Types. The type of Testudo ferox is still extant in the British
Museum.1 The species was discovered, described and illustrated by an
American; his descriptions and illustrations were published by one of
the most outstanding English zoologists of the period in the foremost
scientific journal of Europe; Linnaeus himself was simultaneously
notified of its discovery. Yet, for twelve years it remained without a
systematic name, until a German bestowed on it the name by which it
is specifically known. This unusual nomenclatorial history and the
inaccessibility of the original description to most zoologists justify its
reproduction here with a brief account of the related circumstances.

Dr. Alexander Garden, of Charleston, who was an ardent student of

1 According to Shaw (Gen. Zool., 3, pt. 1, 1802), the specimen was already then in the British
Museum. In 1831 Gray (Syn. Rept., p. 46) confirms that statement and adds that Pennant
gave the type specimen to the Royal Society. In 1844 and 1856 he records it as the "Specimen
described and figured by Pennant, (restuffed)."

28 bulletin: museum of comparative zoology

the natural history of the Carolinas, became possessed of an adult
specimen of the first softshell turtle reported from America. On
December 24, 1770, he wrote to his friend Mr. John Ellis in London
as follows:1

"I have one or two things which I think will please him [Thomas Pen-
nant] ; he shall have them by one of our spring ships ; one of them is a
species of Turtle, as yet nondescript. It is amazing how Catesby
omitted this. It is found in abundance in the Savannah river, in
Altamaha, and East Florida. It is a fresh water animal, grows to a
great size, and is as delicate as the Green Turtle, having a large
leathery cover over its back, and a head very like a Mole. I intend to
send a copy of my account of this animal to Mr. Pennant for his
American Zoology, and if I can get a drawing of it copied, I will send
him that. If I can obtain another Turtle, I shall send you one stuffed.
It has a relation to the first species of Linnaeus's last edition of the
Systema Naturae."2

Early next spring he kept his promise and sent Pennant the descrip-
tion as well as a preserved specimen. Another, better specimen, he
forwarded at the same time to Mr. Ellis with a drawing made from
life. This specimen with the drawing and the letter Pennant laid be-
fore the Royal Society in London on May 2, 1771, in the Philosophical
Transactions of which, vol. 61, they were published. Dr. Garden does
not state exactly whence the specimens came. In his letter he says that
"they are not commonly got here in Charles-town, though by chance
this last summer, I had two sent me", but as he mentions Savannah
and Altamaha rivers, it may be inferred that the two turtles were sent
him from one of those rivers, or one from each of them. One of the
specimens, "the specimen described and figured by Pennant (re-
stuffed)", according to Gray is the type in British Museum.

Dr. Garden's original description follows :

[p. 268] "They are found in large quantities in Savannah and Alta-
maha rivers; and I have been told that they are very common in the
rivers in East Florida.

They grow to very large sizes, though the largest that ever I heard
of was seventy pounds.

The Turtle, which I now have by me, weighs twenty pounds; and
probably, when I first got it, it might have weighed from twenty-five
to thirty pounds, as I have observed that it has grown poorer every

1 Smith, J. E. : A Selection of the Correspondence of Linnaeus and other Naturalists. London,
1821, 1, p. 580.

2 Linnaeus, Systema Naturae, ed. 12, 1, 1766, p. 350 (Testudo coriar.en\.

stejneger: American soft-shell turtles 29

day. I have had it now near three months, and I never could observe
that it has eaten any thing that has been given it, though a variety of
things have been tried.

It is twenty inches long from one end of the shell or covering to the
other, and fourteen inches and a half bread. The colour of this shell
or covering, in general, is dark brown, with a greenish cast.

The middle part is hard, strong, and bony; but all round the sides,
especially towards the tail and hindermost part, it is cartilaginous,
soft and pliable, resembling thick tanned sole-leather, yielding very
easily to any force in any direction whatever, but thick enough and
strong enough to defend the animal from any injury. All the hind
part of the back is full of oblong smooth knobs; and the fore part,
just where it covers the head and neck, is studded full of large knobs.
The under side of this plate is very [p. 269] beautiful, of a lively whitish
colour, interspersed with innumerable very fine ramifications of blood
vessels, running from the margin of the plate into larger and larger
branches, until the sight of them is at once lost by their entering the
body of the animal.

The under, or belly plate, or rather sternum, is of a fair whitish
colour, and extended forward two or three inches more than the back
plate, so that the head rests on it very conveniently. The hind part
of this plate is hard and bony, shaped very much like a man's riding
saddle, with two pieces for the thighs to rest on. The fore part of the
plate is pliable and cartilaginous.

The head is somewhat triangular and attenuated, rather apparently
small for the animal, but growing gradually larger towards the neck,
which is thick and long, and easily extended out (neck of the present
subject was thirteen inches and a half long) to a great length, or drawn
back again under the shell or plate.

The eyes are placed in the fore and upper part of the head, near to
one another, having pretty large loose palpebrae. The pupil is small
and lively, surrounded by a lemon-coloured iris, perfectly round, and
giving much life and fire to the eyes. When danger approaches, or
when it goes to sleep, it covers its eyes, by bringing the inner and
loose part of the lower palpebrae over its eye, like a membrana nictitans.

The upper lip and under lip are both large, but especially the upper.
The mandibula are both entire, each being one entire bone all round,
of the same shape as the mouth.

[p. 270] The nostrils are the most singular part, being a cartilagi-
nous production of at least three quarters of an inch, beyond the upper
and fore angle or point of the upper lip, perforated with two apertures

30 bulletin: museum of comparative zoology

reaching back and opening into the roof of its mouth, having a smooth
septum but fimbriated upon each side. This, at first sight, in some
manner resembles the snout of the mole; but it is tender, thin and
transparent, and cannot be intended for digging in the earth or land.

The arms are thick and strong, consisting of three distinct joints,
viz. the upper, the fore arm, and hand. The hands have each five
fingers, of which the three first are shorter and stronger, and furnished
with strong nails, or rather claws. The two last fingers have more
joints, but are smaller, and, instead of being furnished with claws, are
covered with the membrane, which is extended even beyond their ex-
tremities. Towards the back or hind part, there are two spurious
fingers, which just serve to support the membrane when extended.
The upper side of these arms and hands are covered with a wrinkled
loose skin, of a dusky greenish colour. The legs consist of the same
number of joints, and have the same number of toes as there are fingers
on the fore-feet, and these are furnished with nails in the same manner,
only there is but one spurious toe. Both the fore and hind legs are
thick, strong, and muscular; and as the animal is very fierce, when it is
attacked or disturbed, it often raises itself on its legs, and will leap
forward to bite its disturber or enemy, which it does with great fury
and violence.

[p. 271] They are likewise very strong, and of a lively whitish
colour, because they are generally, if not always, covered with the
upper plate, which, as I said before, is extended a great way behind.

The tail is large and thick, and generally as long as the hind part of
the upper plate. The anus is placed about an inch from the extremity
of the tail on the inside.

The turtle, from which these characters were taken, was a female;
after she came into my possession she laid fifteen eggs, and about the
same number were taken out of the belly when she died. The eggs were
nearly an inch diameter, and perfectly spherical.

It is esteemed very good eating, and said by many to be more
delicate than the green turtle."

On June 20, 1771, Dr. Garden wrote a letter in Latin to Linnaeus
which is translated as follows :l

"I have described and have lately sent to our friends Ellis and

Pennant, a new and very rare species of river Tcstudo, known here

by the name of the Softshelled Turtle, because the covering of its

back, especially towards the sides, is of a softish, leathery, very

1 Smith, op. cit. p. 336.

stejneger: American soft-shell turtles 31

flexible substance. This animal is found in the larger fresh-water
rivers of East Florida, Georgia and South Carolina."
Dr. Garden probably was hoping that Linnaeus might have hastened
to supply the still missing nomcn triviale as he did five years earlier
with Garden's no less startling discovery of Siren lacertina.

It will be noted that Dr. Garden himself, in the letter to Linnaeus,
identified the new discovery with the softshell occurring in East
Florida, and, subsequently, naturalists applied the name ferox indis-
criminately to all specimens from North America. Therefore, when
Lesueur in 1827 described T. spinifcrus as a distinct species and his
great countrymen Cuvier and Dumeril declared it to be only the young
of T.ferox, the identity of the two names was generally accepted, even
by Holbrook (1842) and Gray (1856). * Not until 1857 when Agassiz
demonstrated the distinctness of spinifcrus did it become generally
recognized, although even Boulenger's treatment (1889) shows that
the true characters of T. ferox, as represented by the type specimen,
were not completely understood.

The type specimen in British Museum was examined by Dr. Georg
Baur on September 7, 1888, as closely as the circumstances then per-
mitted. He noted particularly its "sehr rauhe Ornament" (very rough
sculpture), and "Keine Spines, sondern Tuberkel" (no spines, but
tubercles) evidently as compared with spinifcrus; "Schadel vom Typus
des Exempl. von Lucas. Vorderer Theil stark beschadigt, stark vorn
abfallend. Unterkief. ganz von jenem Typus. Jugale mit minim, unt.
Fortsaz." (Skull of the type of Lucas' specimen. Front part much
injured, greatly inclining anteriorly. Lower jaw entirely of that type.
Jugal with minimal lower process). He sums up thus: "Resultat : Type
von Platy pcltis ferox Exemplar von Lucas."

Lucas' "exemplar" is U.S.N.M. 8899 (tintag read upside down 6688
by Baur in his notes) which Baur had been studying with F. A. Lucas,
then curator of the division of comparative anatomy in the National
Museum. No. 8899 is a fine disarticulated skeleton still in the Museum,
of approximately the same size as Dr. Garden's type, and was col-
lected by Professor S. F. Baird, April 1877, in the St. John's River,
Florida, probably not far from Jacksonville. A fine plaster cast
painted by Schindler from his color sketch of the living specimen is
on exhibition in the Museum. A photograph of it was published in
the report of the Museum for 1882, pi. 32.

1 For some unexplained reason, however, Dumeril and Bibron ignored the priority rights of
ferox and called the combination which included the South Carolina to Florida records Gymnopus
iferus. Possibly as a disguised protest against Geoffroy's inefficient substitution of Trionyx
for Schweigger's earlier Emyda.

bulletin: museum of comparative zoology

Thanks to the authorities of the British Museum I have been per-
mitted to examine the type and can confirm completely Dr. Baur's
result as demonstrated by the photographs side by side (pi. 5) of the
two specimens and the measurements in table 1.

Type of Trionyx harlani. In the Medical and Physical Researches
published by him in 1835 Dr. Richard Harlan includes on p. 159 the
description of a Trionyx as follows :
"Trionyx Harlani
Trionyx Harlani, Bell, Monogr. Test, pi.
Char. Body more ventricose, soft portions of the shell less exten-
sive than in the other species. In general appearance ap-
proaching more to the genus Emys.

Inhabits East Florida. Mus. of Bell, Lond."
I have not been able to locate any such plate in Bell's Monograph
of the Testudinata. Bell, in correspondence, may have indicated that
he had in his collection a specimen from East Florida which he intended
to figure in his unfinished Monograph under the above name, but there
is no such plate among the unedited ones published in 1872 under the
title: "Tortoises, Terrapins and Turtles", neither have I found any
record of what became of Bell's specimen. J. E. Gray in the introduc-
tion of his catalogue of the Tortoises in British Museum, notices that
"the specimens presented by .... Thomas Bell, Esq. [may be re-
garded] as the types of the species described in his various papers, and

in his very beautiful Monograph of the Testudinata [by] Dr.

Richard Harlan, and Messrs. Edward and Henry Doubleday, as the
types of the North-American species described by Say, Harlan, and
others," but he has no reference to any softshell from Florida, nor is
there in any of the later catalogs.

External characters. While the skull characters are only available
in dubious and critical cases, normal specimens of A. fcrox within the
group are not difficult to identify, the older ones by the coarseness of
the sculpture of the bony carapace; the very young ones by their
unique coloration.

The coarseness of the sculpture of the bony carapace of A. ferox is
not confined to the general network of vermiculations of the surface,
but it is commonly specialized into a series of more or less prominent
longitudinal welts. The difference in relative size and pattern of the
pits and ridges which constitute the character of the "sculpture" is
difficult to describe, but a comparison of the samples figured (pis. 7,
9, 10) explains it better than words.

The unique coloration of the hatchlings and the very young speci-

stejneger: American soft-shell turtles

mens constitutes the most obvious and characteristic feature of the
species, but unfortunately disappears with age. In all the other species
the young are of a more or less pale olive or tawny ground color, either
uniform or marked with dusky or blackish specks of varying shape,
but mostly round dots, which when larger assume the form of ocelli
with a lighter center, combined with a nearly uniform white plastron
(except for the pinkish tinge due to the fine blood vessels shining
through in the living specimens). The prevailing feature is the dark-
ness and the saturation of the pigmentation, the big dark blotches on
the carapace, and the dark slate gray underside, in combination with
the strong contrast of the light pattern on the head and the margin of
the carapace, detailed description of which will be given below.

The anterior flap of the leathery plastron is longer and less circular
than in the other species, often extending a considerable distance for-
ward beyond that of carapace, but it appears to vary individually and
is difficult of precise definition because of absence of suitably fixed
points from which to measure.

The shape of the outline of the disk, however, is somewhat different
in the }roung of A. ferox, in as much as it is less circular than in the
other species, as will appear from table 3 which presents the measure-
ments of five hatohlings of the same brood. Similar proportions are
shown by one collected by Dr. Francis Harper in the Okefinokee
Swamp (U.S.N.M. 84603), viz. length of leathery carapace 39.5 mm.
and width 32.5 mm. It is also well illustrated in the photograph of
No. 61087, pi. 17, fig. a.

Table 3

Amyda ferox, pullus

Florida: Polk Co., Auburndale

mm.

Length of leathery disk
Width of leathery disk mm.
Height of body

mm.

61083

38.0
33.5

12.5

61084

40.5
35.5
12.0

61085

40.0
34.0
12.5

61086

41.0
34.0
12.0

61087

45.0
37.0
14.0

Average

40.9
34.8
12.6

Size. Amyda ferox is apparently the largest of the North American
softshell turtles. Authentic measurements of large carapaces are few

34 bulletin: museum of comparative zoology

and those available are of dubious value, due to uncertainty as to
identification, condition of specimen when measured, and method of
measurement whether in a straight line or along the curvature of the
shell. The largest Agassiz "had ever seen or heard of" was one from
Natchez "which measured eighteen inches and a half [470 mm.] from
the front to the hind margin of the carapace" (Contrib. Nat. Hist.
U. S., vol. 1, p. 401). The largest specimen now in the National
Museum is an old skin, with the skull in (U.S.N.M. No. 38123 from
"Florida" which measures about 17 inches along the curvature (430
mm.). Its zygomatic width is 68 mm. The plaster cast of No. 8899 —
(in exhibition series) is 438 mm. long; over the curvature it measures
460 mm. (18 inches); zygomatic width 67 mm.

But larger specimens may exist, or have been living in Florida not
long ago. The National Museum has a series of 16 weathered skulls
picked up by Dr. E. A. Mearns near Kissimee about the beginning of
this century, 14 of them larger than that of the 18 inch specimen men-
tioned above (No. 8899). The basicranial length of this one is 92 mm.,
the corresponding dimension of the 14 Kissimee skulls range from 95
to 114 mm. (aver. 103) with the zygomatic width varjing between
65 and 80 mm. (aver. 73).

Abnormal alveolar surfaces

This series of 16 skulls of evidently very old specimens shows an
extraordinary development which deserves special attention.

In the normal skulls of the species in all our specimens with a basi-
cranial length below 90 mm. the lateral outline of the snout anterior
to the orbit tapers towards the end in a fairly straight line, and the
narrow maxillary alveolar surface follows almost parallel, as in the
other species of the ferox group, irrespective of sex and age.

However, in the series of 16 Mearns skulls with basicranial length
above 84 mm. we find two different styles of snout outline and alveolar
surface as recorded in table 4.

For the illustration of the extremes of the width of the alveolar sur-
face and the outline of the maxilla it is only necessary to refer to plate
6. While greatly reduced the figures convey the difference between the
two series as the figures are of the same relative size.

Evidently the abnormal development of the maxilla as represented
by plate 6, figs. 3 and 4 are due to old age since we do not find it in the
smaller and younger specimens. It at once recalls similar conditions
in some Chinese species of the genus, upon which Father Heude, in

stejneger: American soft-shell turtles

Table 4
Alveolar width and outline of snout in a series of very old A. ferox.

Snout outline straight

Snout outline strongly

convex

Greatest

U.S.N.M.

alveolar

Basicranial

alveolar

U.S.N.M.

No.

width

length

width

No.

029459

7.5 mm.

114 mm.

108 mm.

21.5 mm.

029464

107 mm.

15.5 mm.

029458

106 mm.

14.0 mm.

029454

105 mm.

14.0 mm.

029470

105 mm.

16.0 mm.

029457

104 mm.

18.0 mm.

029463

104 mm.

11.5 mm.

029451

101 mm.

15.0 mm.

029450

101 mm.

11.5 mm.

029460

100 mm.

14.0 mm.

029455

029456

7.5 mm.

100 mm.

97 mm.

11.5 mm.

029453

95 mm.

9.5 mm.

029452

029475

6.8 mm.

92 mm.

84 mm.

12.0 mm.

029462

1880, 1 based the description of numerous new genera and species. I
need only refer to his pictures of Caelognathus novemcostatus, pi. 5;
Tortisternum novemcostatum, pi. G; Cinctisternum bicinctum, pi. 9; and
especially Ceramopelta latirostris, pi. 7. Notwithstanding the fact that
he did not obtain any young specimens with these characteristics
Heude regarded them as specific or generic differences. Boulenger in
cataloguing the softshell turtles in the British Museum (1889, p. 243)
was "unable to find a single young specimen with the molar-
like alveolar surfaces" but having "found in three species, viz. T.
triunguis (Africa), T. cartilagineus (E. Indies), and T. sinensis (China)

i Mem. Hist. Nat. Emp. Chinois, 1, pp. 1-38, pis. 1-9.

36 bulletin: museum of comparative zoology

examples of the two types, i.e. on the one hand sharp-edged, com-
paratively narrow jaws, and on the other hand broad crushing alveolar
surfaces nearly meeting on the median line in front of the choanae,

in specimens which, in other respects, are undistinguishable,

[he] arrived at the conclusion that we may be in presence of a case of
dimorphism caused by a difference of diet."

This idea of a "dimorphism" due to a difference of diet does not seem
convincing. That it is not the result of old age alone seems obvious
from the fact that so many old skulls of the same size and presumably
age do not show the abnormality. That it is not due to a change of
diet affecting the whole population seems also obvious since the Kissi-
mee series are all practically from the same locality. That the differ-
ence is not a local one is proved by a weathered skull almost identical
with Kissimee No. 029462 of the above table, picked up by Dr. Francis
Harper in the Okefinokee Swamp (U.S.N.M. No. 59727) which with
a basicranial length of 88 mm. has a maxillary alveolar width of 11
mm. The fact that in the series the largest skull is normal and the
smallest abnormal coupled with the other fact that all the largest
authentically sexed specimens in the collection are female and normal,
while among the preserved specimens no abnormal male has been
recorded, suggest that the difference may be due to sex. If so, does that
indicate an individual preference among the old males for a certain
kind of diet?

The normal skull. As repeatedly noted the skull of A. ferox is built
on the same plan as that of spinifera and cmoryi and differs but slightly
in the proportionate size and relation of the various bones, but the
individual variation is so great that in some cases it is even difficult
to decide to which species an isolated skull without locality record be-
longs. A. ferox, however, may generally be diagnosed as having the
narrowest interocular width, the shortest distance between choanae
and intermaxillary foramen, and, in the medium-sized skulls, the
narrowest maxillary alveolar surface. It also averages the greatest
length from orbit to tympanic cavity combined with the shortest and
weakest mandibular symphysis. The interorbital space is usually less
in width than one third of the longest diameter of the temporal fossa,
while in the others it is usually more than one third. The anterior
processes of the prefrontals as a rule are longer and slenderer and the
angle projecting into the posterior border of the nasal fossa is conse-
quently more acute. In the older and younger specimens these differ-
ences become more obscure or may be entirely obliterated.

In a series of skulls of this group a rather common feature may be

stejneger: American soft-shell turtles 37

noted in the larger specimens of A. ferox. While in the palate of the
other species the maxillaries are in contact practically the whole length
between the choanae and the intermaxillary foramen, they may be en-
tirely or partly separated by the vomer which in many skulls may be
seen as a fork enclosing the posterior end of the foramen as shown in
all the figures on pi. 6 (very large specimens).

Coloration. Old specimens of this, as well as the other species, show
but little of the characteristic normal coloration and pattern of the
species. In A. ferox more or less faint remnants of the large brownish
blotches on the carapace may be made out on the generally dingy
"Isabella" colored ground, or disappear in the blackish or brownish
variations of the latter. But as previously indicated, the very young
ones display a distinctive and peculiar color and pattern, unknown
until figured by Ditmars (1907) and described by Wright and Funk-
houser (1915). Some of the specimens received by the National
Museum were either alive or freshly killed and their descriptions with
reference to a standard color nomenclature (Ridgway's Nomenclature
of Colors for Naturalists, 1886) were made:

U.S.N.M. No. 61087. Auburndale, Polk Co., Florida, collected by
N. R. Wood, summer of 1918. Pullus. Length of leathery carapace,
45 mm. Leathery plastron extending anteriorly beyond carapace 6 mm.
Iris pale silver gray with a horizontal black bar. — Upper surface of
carapace tawny olive (Ridgway, pi. iii, fig. 17) with dusky (dark
sepia) spots and a narrow well-defined outer edge of bright ochraceous
(R. v, 7) in strong contrast; underside of carapace and plastron slate-
gray (R. ii, 5) with scattered clay-colored spots on the former and the
anterior edge of plastron, the outer edge of disk narrowly ochraceous,
as above, though less bright; upper side of head, neck and legs olive
(R. iii, 9) with clay-colored marblings; from anterior angle of eyes to
middle of proboscis an inverted Y-shaped, pale clay-colored figure, the
fork situated halfway between eye and base of proboscis; side of head
olive with a wide well-defined angular band of yellowish buff extending
from posterior corner of eye to base of lower jaw; another similar
band, but slightly broken and brighter, almost cadmium-orange
(R. vi, 2) anteriorly, originating behind the former and descending on
the neck to past the middle; a third band, paler buff, curving around
the corner of the mouth almost meeting below the corresponding band
of the other side on the posterior third of the neck; a fourth, median
band between the last ones; soft skin of feet almost "plumbeous"
(R. ii, 15) underneath.

Four other specimens of the same brood, Nos. 61083-86, are colored

38 bulletin: museum of comparative zoology

essentially as the above. The numbers, sizes, shape and arrangement
of the dark blotches on the carapace vary to a great extent and so does
consequently the light network (really the ground color of the cara-
pace) separating them, it being slightly wider in the specimen figured
(pi. 19, fig. a). There is one feature common to all and of some signifi-
cance, viz. the more or less parallel arrangement of the outer blotches
on the posterior flap, those of the outer row nearly forming a continu-
ous dark line, the next row also coalescing on the posterior half, thus
clearly indicating the blackish submarginal rings of the other species,
as shown in the illustration just quoted.

Wright and Funkliouser (Proc. Acad. Nat. Sci. Philadelphia, vol. 67,
1915, p. 122) note that "as the specimens become older, the gayly
colored markings of the carapace become less distinct and have dis-
appeared on turtles which have attained a length of 6 inches [152 mm.]."
The National Museum has a specimen (No. 56804) from Irwin Co.,
Georgia, the carapace length of which is 100 mm., which still shows the
pattern as described above, but the tubercles on the disk are all well
developed.

Their further statement that with age "the plastron grows lighter in
color and the head uniformly darker with the markings obsolete", is
borne out by the following description made in July 1919 of a live male
specimen (now U.S.N.M. No. 62217) from Georgia, Berrien County,
with a leathery disk about 235 mm. long and 195 mm. wide: Iris
brownish gray with a brassy edge against the pupil. Carapace dark
"raw umber" (R. iii, fig. 14) with large dusky, more or less confluent
irregular marblings; top of head, neck and legs "sepia" (R. iii, fig. 3)
with paler marblings ; proboscis, snout, sides of head, and lips strongly
washed with "cinnamon" (R. iii, fig. 20); a dark brown mark across
the forehead in front of eyes continued behind them to the ear as a
series of spots, and a few spots of the same color behind corner of
mouth; on underside of neck back of the skull three longitudinal white,
dusky -edged marks; underside of body white; digits and webs strongly
washed with plumbeous (R. ii, fig. 15).

Some larger specimens may still retain traces of the original pattern.
A full grown male (U.S.N.M. No. 60496, carapace length 316 mm.)
collected at Auburndale, Florida, was received alive on February 28,
1918, and its colors at once described by me as follows: Iris dark silvery
gray with a darker horizontal bar and the inner edge forming a com-
plete bright narrow silvery ring sharply defining the pupil; inside of
nostrils dark pink; inside of mouth pale flesh-color. Ground color of
carapace bistre with anastomozing, ill-defined lines (10 to 15 mm. wide)

stejneger: American soft-shell turtles 39

of tawny olive, which isolate islands of the ground color of varying
sizes but averaging perhaps 25 mm. in diameter; neck and legs above
dark olive, head anteriorly suffused with cinnamon changing to pink
on proboscis; underside pinkish white, on neck suffused with dark
purplish gray on which dusky-edged cream-colored lines here and there
with touches of orange; palms and soles dark olive.

Aberrant Color Phase. A color anomaly shown by a specimen, the
history and description of which, because of the uniqueness of the pig-
mentation deserves to be recorded, is in the National Museum.

On April 11, 1881, there was entered in the Museum register of
reptiles a consignment of living specimens collected in March by James
Bell at Gainesville, Florida. Among them was an "Aspidonectes" of
which the artist Z. Schindler on April 9 had made a water-color sketch
for use in later painting the plaster cast for the "exhibition series".
The specimen, a female (Xo. 10545, carapace 225 mm. long, 16S mm.
wide), now a skin with skull separate, the color sketch, and the painted
plaster cast (slightly broken) are still in the Museum. The water-color
sketch and the oil-painted cast agree in all essentials, except that the
cast is (now) considerably darker. The skull and other structural
characters are those of a normal Amydaferox, but the color is an ex-
treme case of erythrochroism. The ground color of the carapace in
the water-color agrees closely with what Ridgway calls "mummy
brown" (R. iii, fig. 10), while in the cast it is more like his "chestnut"
(R. iv, fig. 9); the throat and front of neck are "rufous" (R. iv, fig. 7)
in both, though lighter in the former; top of head, back of neck, and
upper side of legs raw umber (R. iii, fig. 14) in the water-color sketch
while in the cast they are dark sepia washed with rufous; the most
conspicuous feature is the band from beneath the eye on the auricular
and temporal region which is bright vermilion in both paintings-
reminding one of the corresponding band in Pseudemys elegans) and
edged with a narrow dusky line anteriorly above and below. The un-
derside, judging from the color sketch (and the specimen) was uniform
white.

Geographical distribution

Florida north to South Carolina and west along the Gulf Coast to
Louisiana.

bulletin: museum of comparative zoology

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bulletin: museum of comparative zoology

List of specimens in U. S. National Museum

4373

Fla.

, Palatka

T. Glover

7651

8708

Ga.

Milledgeville

T. H. Bean

July, 1876

8899 9 ad.

St. Johns R.

S. F. Baird

Apr. 3, 1877

9670 d1 adol.

S. C

!., Charleston

C. C. Leslie

May, 1878

10545 9 adol.

Fla.

, Gainesville

J. Bell

Mar., 1881

10704

19621 adol.

Ga.

Darien

20189 adol.

Fla.

, Eustis

T. Holm

March, 1893

26035 juv.

ponds near

Welaka

W. C. Kendall

Mar. 20, 1897

29210 9 ad.

E. A. Mearns

029339 adol.

Eustis

H. J. Webber

029448-9

Kissimmee

E. A. Mearns

029450-1

029452-9 c? ad.

029460-2 c? ad.

029463 tf1 ad.

029464-6 cf ad.

029467-8

029470 cf ad.

029474 9 ad.

029475 9 ad.

029619 adol.

Ga.,

Mimsville

C. S. Brimley

38123 9 ad.

Fla.

M. L. Odell

38980-1 juv.

Ga.,

Mimsville

C. S. Brimley

June 19, 1909

51184 juv.

Fla.

ponds near

Tampa

Evermann & Kendall

Nov. 3, 1896

51417-20 hf-gr.

St. Petersburg

C. S. Brimley

51421 juv.

Orlando

52476-83

Fla.,

Eureka

C. S. Brimley

Aug. 12, 1915

55316 c* ad.

Fla.

Vero

I. M. Weills

56804 juv.

Ga.

Irwin Co.

J. Hurter

Oct. 15, 1902

56805

Fla.,

Orlando

July, 1911

56806

Hillsboro Co.

June 15, 1910

56807 ad.

Bronson

1891

59318

Sebastian

F. Harper

Feb. 23, 1917

59727-8 (skullk

only) <? ad.

Fla.,

Lake Okeechobee F. Harper

Feb. 6, 1917

60496 c? ad.

[Auburndale]

N. R. Wood

Feb., 1918

60532 adol.

Auburndale

60533 ad.

60534 9 ad.

stejneger: americais

r SOFT-SHELL TURTI

,ES <i«5

60547 & ad.

Fla., Auburndale

N. R. Wood

1918

60556 juv.

« a

■ a

Mar., 1918

60S28 eggs

" Homestead

C. A. M osier

60902-4

" Eureka

C. S. Brimley

1918

61031

" Gulf port

A. G. Reynolds

61083-7 pull.

" Auburndale

N. R. Wood

July 22, 1918

61096-109

ii it

Aug. 24, 1918

61352 9 ad.

" Lake Miakka

C. M. Barrett

June 18, 1918

62217 & adol.

Ga., Banks Mill Pond

U.S. Bur. Fish.

63343

Fla., Auburndale

N. R. Wood

1920

70398

Ga., Mimsville

C. R. Brimley

71068-9

71156 adol.

Fla., Plant City

C. R. Aschemeier

1926

71681 !

S.C., Greenwood

Dr. Barrett

73199

Fla., Delray

J. D. Thieme

80963

Fla.

F. H. Benjamin

Apr., 1930

84079

" 15 mi. from

Miami

M. K. Brady

1930

84080

" Orlando

E. T. Evans

84603 pull.

Ga., Chesser's Id.,

Okefinokee Swamp

F. Harper

July 21, 1931

86492 d*

Fla., 15 mi. from

Miami

M. K. Brady

Mar., 1932

86828 <? adol.

" nr. Birdon

P. Bartsch

Aug., 1932

95767 <? adol.

" Lake Iamonia

C. R. Aschemeier

Feb. 16, 1935

103736 Juv.

" Silver Lake

Mar. 7, 1938

Amyda spinifera (Lesueur)

Agassiz, in 1857, while demonstrating the distinctness of Lesueur's
species described 30 years before from the Wabash River against the
contention of contemporary authors that it was only the young of
Amyda ferox, at the same time established three additional species,
Aspidonectes as per from the state of Mississippi, A. nuchalis from
Tennessee, and A. emoryi from Texas. The latter has been generally
accepted, nuchalis has been generally ignored, and asper only recently
recognized as a distinguishable race of A. spinifera. As such it will be
treated below trinominally.

The characters attributed to A. nuchalis have lost their significance
after the accumulation of additional material of A. spinifera, and the
study of available series of specimens in connection with the present
investigation have not yielded data indicating a separable group popu-
lating the upper reaches of the Tennessee and Cumberland Rivers.

'Identity questioned.

44 bulletin: museum of compakative zoology

AMYDA SPINIFERA SPINIFERA1 (LeSUeUr)

Plates 11-15

1825. — Trionyx ferox SAY, Journ. Acad. Nat. Sci., ser. 1, 4, pt. 2, p. 218 (part:
Mississippi, Ohio, and Missouri Rivers; New York, etc.) (not of
Schneider).— LeCONTE, Ann. Lync. Nat. Hist., New York, 3, 1830,
p. 93 (part: all streams which run into the Mississippi). — HARLAN,
Med. Phys. Res., 1835, p. 158 (part).— SCHLEGEL, Fauna Japon.,
Rept., 1838, p. 30, pi. 5, fig. 5 (Cumberland, Tennessee and Ohio
Rivers).— HOLBROOK, North Amer. Herpet., 1 ed., 4, 1840, p. 9
(part); 2 ed., 2, 1842, p. 11, pi. 1 (part: Mississippi; great northern
lakes; Mohawk River, New York). — DeKAY, Zool. New York, pt.
3, Rept., 1842, p. 6, fig. 11 (Mohawk River; Hudson River near
Albany, New York).— TROOST, Seventh Geol. Tennessee, 1844,
p. 39 (Tennessee).— THOMPSON, Hist. Vermont, 1853, p. 29
(Vermont: Rivers Lamoille and Winooski). — GRAY, Cat. Tort.
Brit. Mus., 1844, p. 49 (part); Cat. Shield Rept., 1, 1856, p. 68
(part).— KENNICOTT, Trans. Illinois Agric. Soc, 1, 1855, p. 591
(Illinois: Cook Co., Lake Michigan).— STRAUCH, Mem. Acad.
Sci. St. Petersbourg, ser. 7, 5, no. 7, 1862 (part); 8, no. 13, 1865, p.
122 (part).— DITMARS, Reptile Book, 1907, pi. 27, upper fig. (not
of text).— GADOW, Cambridge Nat. Hist., 8, 1901, p. 408 (part),
p. 409, fig. 92 (North America).
Amyda ferox ORTENBURGER, Copeia, no. 170, May 1929, p. 12
(Oklahoma: LeFlore Co.); p. 28 (Oklahoma: Rogers Co.).

1827 .—Trionyx spiniferus LESUEUR, Mem. Mus. Hist. Nat. Paris, 15, Dec-
1827, p. 258, pi. 6 (type-locality, Wabash River, New Harmony*
Indiana; types in Paris Mus.; Lesueur, collector). — WIED, Reise
Nord-Amerika, 1, pt. 3, 1838, pp. 140, 141 (Pittsburgh, Pa.); Voy.
Amer. Nord, 3, 1843, p. 242.— HAY, Indiana Geol. 17 Rep., 1892, p.
554; Batr. Rept. Indiana, 1893, p. 146 (Indiana: generally distri-
buted).—HURTER, Trans. Acad. Sci. St. Louis, 6, 1892, p. 260
(Missouri: Mississippi River, Merimac River, and Illinois River). —
HAHN, Proc. U. S. Nat. Mus., 35, Dec. 1908, p. 567 (Lawrence Co.,
111.).— SIEBENROCK, Zool. Jahrb. Suppl., 10, pt. 3, 1909, p. 604
(Mississippi River and tributaries; St. Lawrence River; Hudson
River) ; Verh. Zool.-Bot. Ges. Wien, 73, Aug. 1923, p. 186.— THOMP-
SON, Thirteenth Rep. Michigan Acad. Sci., 1911, pp. 106, 107, fig. 1
(Cass Co., Michigan).— ELLIS and HENDERSON, Univ. Colorado
Stud., 10, no. 2, May 1913, p. 112 (Colorado: Weld Co.: Evans,
Cache la Poudre, South Platte River, and Greeley).

1 Spine bearing, with reference to the pointed shape of the tubercles margining the anterior
flap of the leathery carapace, characteristic of older specimens.

stejneger: American soft-shell turtles 45

Gymnopus spiniferus DUMERIL and BIBRON, Erpet. Gen., 2, 1835,
p. 477 (atlas, pi. 22, fig. 1) (Wabash River).— DUMERIL, Cat.
Meth. Coll. Rept. Mus. Hist. Nat. Paris, 1851, p. 22 (Wabash River).
— SAGER, Peninsular Journ. Med. Collat. Sci., 3, no. 8, 1856, p. 361
(anatomy). — WIED, Nova Acta Acad. Leopold. -Carol., 32, pt. 1,
1865, p. 48 (Wabash River).

Gymnopodus spiniferus DUMERIL, Arch. Mus. Hist. Nat., 7, 1856,
p. 203.

Aspidonectes spiniferus RHOADS, Proc. Acad. Nat. Sci. Philadelphia,
1895, p. 386 (Tennessee: Samburg, Obion Co.).

Aspidonectes spinifer AGASSIZ, Contr. Nat. Hist. United States, 1,
1857, p. 403; vol. 2, pi. 6, figs. 1-2 (Lake Champlain; Lake Ontario
and Erie; New York, Pennsylvania, Ohio, Indiana, Illinois, Missouri,
Michigan, Wisconsin, Iowa, Ft. Union, Montana). — MILES, First
Biennial Rep. Geol. Surv. Michigan, 1861, pp. 232, 233 (Michigan:
as far W. as Genessee Co.). — ALLEN, Proc. Boston Soc. Nat. Hist.,
1874, p. 69 (Musselshell and Yellowstone Rivers, Montana). —
COPE, Bull. U. S. Nat. Mus., No. 1, 1875, p. 51.— JORDAN, Man.
Vert. North Amer., ed. 3, 1880, p. 168; ed. 5, 1888, p. 206 (Canada
to Kentucky and Minnesota); ed. 8, 1899, p. 206.— CRAGIN,
Trans. Kansas Acad. Sci., 2, 1881, p. 116 (Kansas: Franklin and
Douglas Cos.).— CONES and YARROW, Bull. U. S. Geol. Geogr.
Surv., 4, 1878, p. 261.— SMITH, Rep. Geol. Survey Ohio, 4, 1882,
p. 668 (all streams flowing into Ohio and Lake Erie). — TRUE, Bull.
U. S. Nat. Mus., No. 24, 1883, p. 29 (Webster City, Iowa; Mt.
Carmel, Illinois; Fox River, Illinois; Ft. Laramie, Nebraska); Fish.
Industr. United States, Sect. 1, 1884, p. 152.— HOY, Geol. Wiscon-
sin, Surv. 1873-1879, 1, 1883, p. 423 (Western Wisconsin).— DAVIS
and RICE, Illinois State Lab. Nat. Hist., Bull. No. 5, 1883, p. 52
(north of Ohio River); Bull. Chicago Acad. Sci., 1, no. 3, 1883, p. 32
(Illinois).— HOY, Geol. Wisconsin, 1, 1883, p. 423 (Wisconsin).-
HUGHES, Bull. Brookville Soc. Nat. Hist., No. 2, 1884, p. 41
(Franklin Co., Indiana).— STOCKWELL, Journ. Comp. Med. Surg.,
9, 1888, p. 28.— HIGLEY, Trans. Wisconsin Acad. Sci., 7, 1889, p.
159 (south and west Wisconsin). — GARMAN, H., Bull. Illinois
State Lab. Nat. Hist., 3, 1892, p. 246 (throughout Illinois: Rock
Creek, Kendall Co.; Piano; Oregon, Ogle Co.; Peoria, Peoria Co.;
Bluff Lake, Union Co.; Wabash River, Mt. Carmel).— KIRSCH,
Bull. U. S. Fish Coram. 1894 (1895), p. 81 (Eel River, Indiana).-
McLAIN, Notes Coll. Rept. Arkansas, 1899, p. 1 (Bloomington,
Monroe Co., Indiana). — SMITH, Proc. Linn. Soc. New York,
1898-99, No. 11, 1899, p. 24.— RAMSEY, Proc. Indiana Acad. Sci.,
1900 (1901), p. 224 (Lake Winona, Indiana).— ATKINSON, Ann.
Carnegie Mus. Pittsburgh, 1, 1901, p. 154 (Monongahela River,

46 bulletin: museum of comparative zoology

Pennsylvania).— MORSE, Ohio Natural., 1, no. 8, 1901, p. 127
(Columbus, Ohio); Proc. Ohio Acad. Sci., 4, pt. 3, Spec. Pap. No. 9,
1904, p. 138 (Ohio: Columbus; Sandusky; London).— PAULMIER,
New York State Mus. Bull. 51, 1902, p. 392 (Lakes Ontario and
Erie; through Erie Canal to Hudson River). — CLARK, Rep. 4
Michigan Acad. Sci., 1904, p. 193 (Eaton Co., Michigan).— JOR-
DAN, Man. Vert. Anim. U. S., 1904, p. 206.— GIBBS, NOTE-
STEIN and CLARK, Rep. 7, Michigan Acad. Sci., 1905, p.
110 (Michigan: Brookfield, Ann Arbor, Olivet, Kalamazoo, Van
Buren, Montcalm and Allegan Cos.). — STONE, Amer. Natural.,
40, 1906, p. 168 (into Delaware Valley; Cooper's Creek and Warren
Co., New Jersey; Allegheny River, western Pennsylvania). — NASH,
Check List Vertebr. Ontario, 1906, p. 17 (Canada: Ontario, western
part; one record Ottawa River). — FOWLER, Rep. New Jersey State
Mus., 1906 (1907), p. 211, pi. 57 (Cooper's Creek; Paulin's Kill,
Hainesburg, Warren Co., New Jersey, introduced). — SURFACE,
Zool. Bull. Pennsylvania Dep. Agric, 6, nos. 4-5, 1 Sept., 1908, p.
121, fig. 2 (Pennsylvania: Indiana Co.; Somerset Co.). — REED and
WRIGHT, Proc. Amer. Philos. Soc, 48, 1909, p. 408 (Cayuga Lake,
New York).
Trionyx spinifer BOULENGER, Cat. Chel. Brit. Mus., 1889, p. 259
(Foxbury, Pennsylvania; Wabash River). — SCHNEE, Zeitschr.
Naturwiss., 72, pt. 3, Dec. 24, 1899, p. 197.— SIEBENROCK, Sitz.
Ber. Akad. Wiss. Wien, Math.-Nat, KL, 111, 1902, p. 829, fig. 10
(plastron).— HENSH AW, Occas. Pap. Boston Soc. Nat. His., 7,
pi. 1, 1904, p. 4 (Lake Champlain, Vermont).— DITMARS, Rep-
tile Book, 1907, p. 77, pi. 26, upper and middle figs.; pi. 28 (Quincy,
Adams Co., Illinois).

Platypeltis spinifer BAUR, Proc. Amer. Philos. Soc, 31, 1893, p. 220
(Wabash River, Indiana).— EVERM ANN and CLARK, Proc.
Indiana Acad. Sci., 1916 (1918), p. 473 (Lake Maxinkuckee, Indiana).

Platyrettis KIRSCH, Bull. U. S. Fish Comm. 1894 (1895), p. 333
(Maumee River Basin, Ohio and Indiana).

Amyda spinifer -POTTER, Copeia, No. 82, Oct. 15, 1921, p. 76 (Thorn-
apple River, Barry Co., Michigan).— SOLA, Bull. New York Zool.
Soc, 34, no. 5, Sept.-Oct,, 1931, pp. 134, 141, 155, fig. 6 (Lake Cham-
plain).

Callinia spinifera GRAY, Proc Zool. Soc. London, 1869, p. 222
{spicifera err. typogr.); 1873, (p. 60, figs.); Suppl. Cat. Shield. Rept.
Brit, Mus., pt. 1, 1870, p. 109.

Amyda spinifera HURTER, Trans. Acad. Sci. St. Louis, 20, 1911, p.
251 (Missouri: Mississippi, Missouri, Osage, Gasconade, Meramee
and White Rivers).— STEJNEGER and BARBOUR, Check List

stejneger: American soft-shell turtles 47

North Amer. Amph. Rept., Ed. 1, Dec. 12, 1917, p. 125 (Mississippi
River and tributaries west to Colorado, north to Montana; St.
Lawrence River and tributaries; east to Vermont, western New
York and Pennsylvania); Ed. 2, 1923, p. 141; Ed. 3., 1933, p. 153.-
WRIGHT, Copeia, No. 66, Feb. 25, 1919, p. 8 (Bays of Lake On-
tario, New York).— BABCOCK, Mem. Boston Soc. Nat. Hist., 8,
no. 3, April 1919, p. 419 (east shore, Lake Champlain, Vermont). —
CLARK and SOUTHALL, Rep. U. S. Comm. Fish 1919 (1920),
App. no. 7, p. 15, pis. 7-8 (economics).— EVERMANN and CLARK,
Lake Maxinkuckee, 1920, p. 592 (habits).— ORTENBURGER,
Copeia, No. 99, Oct. 15, 1921, p. 76 (Decatur Co., Indiana); Proc.
Oklahoma Acad. Sci., 6, pt. 1, 1926, p. 100 (McCurtrin and Push-
nataha Cos., Oklahoma). — BLANCHARD, Occas. Pap. Zool. Mus.
Univ. Michigan, No. 117, July 6, 1922, p. 18 (Reelfoot Lake, West.
Tennessee); Univ. Iowa Studies Nat. Hist., 12, no. 2, 1923, p. 24
(Little Sioux Riv., Dickinson Co., Iowa); Pap. Michigan Acad. Sci.,
5, 1925, p. 386 (10 miles W of Columbus, Indiana).— WEED,
Copeia, No. 116, March 15, 1923, p. 48 (Meredosia, Illinois).—
BISHOP, Copeia, No. 125, Dec. 31, 1923, p. 120 (Albany Co., New
York).— PRATT, Man. Vert. United States, 1923, p. 249.— LO-
GIER, Canad. Field Natural., 39, May, 1925, p. 95 (Point Pel6e;
Hamilton, Dundas Marsh, Ontario, Canada); Roy. Ontario Zool.
Mus. Handb. no. 4, 1939, p. 56, pi. 7, fig. 2; pi. 8, fig. 7 (Ontario:
Hamilton Bay and Dundas Marsh, Wentworth Co.; Thames River
at Beachville, Oxford Co.; Grand River at Dunnville, Haldimand
Co.; Long Point, Norfolk Co., and Point Pelee, Essex Co.). —
SCHMIDT, Copeia, no. 154, May 20, 1926, p. 132 (Chippewa Co.,
Wise.).— MYERS, Proc. Indiana Acad. Sci., 35, 1926, p. 292 (In-
diana); vol. 36, 1927, p. 339 (Helmsburg, Indiana). — BURT, Occas.
Pap. Zool. Mus. Univ. Michigan, no. 189, Dec. 12, 1927, p. 9 (Riley
Co., Kansas).— LINSDALE, Copeia, no. 164, July-Sep., 1927, p. 81
(Doniphan Co., Kansas); Trans. Kansas Acad. Sci., 36, 1933, p. 208
(Cowley and Lane Cos., Kansas).— RUTH VEN, THOMPSON, and
GAGE, Herpet. Michigan, 1928, p. 163, pi. 19, fig. 3 (Michigan).-
MERTENS, Zool. Gart. Leipzig, new series, 1, pt. 5-6, 1928, p. 199
(distr. New York State by canals). — GLOYD, Trans. Kansas Acad.
Sci., 31, 1928, p. 135 (Franklin Co., Kansas).— POPE and DICKIN-
SON, Bull. Publ. Mus. Milwaukee, 8, no. 1, Apr. 3, 1928, p. 82, pi. 21,
figs. 7-8 (Wisconsin: Burnett, Crawford, Grand, Oneida, Polk,
Waukesha, Chippewa, Washburn, and Pepin Cos.). — POPE, Year
Book Mus. Milwaukee, 1928 (1929), pp. 180, 183 (Vernon and
LaCrosse Cos., Wisconsin). — JORDAN, Man. Vert. Northeast U.S.
(13 ed.) 1929, p. 254.— CAHN, Copeia, no. 170, May 1929, p. 8
(Lake La Belle, Waukesha Co., Wisconsin). — FORCE, Copeia,
1930, no. 2, p. 38 (Tulsa Co., Oklahoma).— DOLLEY, Amer. Mid-

48 bulletin: museum of comparative zoology

land Natural., 14, no. 3, May, 1933, p. 203 (St. Joseph Riv., Berrien
Co., Michigan).— RUST, Blatt. Aquar. Terrarienk., 45, 1934, p. — ,
sep. p. 12.— BURT and HOYLE, Trans. Kansas Acad. Sci., 37,

1934, p. 198 (Kansas).— BURT, Amer. Midland Natural., 16, no. 3,

1935, p. 321 (Kansas: Barber, Reno and Sedgwick Cos.).— BOYER
and HEINZE, Trans. Acad. Sci. St. Louis, 26, no. 4, Apr. 1, 1934, p.
199 (Missouri: Jefferson Co., common).— NETTING, Nawakwa
Fireside (N.S.), nos. 3-4, Apr., 1935, p. 49 (Penna,: 8 counties);
Proc. Pennsylvania Acad. Sci., 10, 1936, p. 27 (Pennsylvania, In-
diana Co., Plum Creek and Crooked Creek).— BRINN, Rep. Penn-
sylvania Fish Comm. for 1938-1939, p. 127 (Pennsylvania: rec. from
9 counties in Ohio and Lake Erie drainages). — SCHMIDT and
NECKER, Bull. Chicago Acad. Sci., 5, no. 4, Sept. 27, 1935, p. 76
(Illinois: Cook Co., Kankakee Co.; Indiana: Lake Co.).— TAYLOR
Univ. Kansas Sci. Bull., 22, no. 11, Apr. 15, 1935, p. 217 (Arkansas:
Devall Bluff, Prairie Co.; Lewisville, Lafayette Co.).— BABBITT,
Bull. Boston Soc. Nat. Hist., no. 78, Jan. 1936, p. 10 (Lake Cham-
plain, Vermont, rare). — HIBBARD, Trans. Kansas Acad. Sci., 39,

1936, p. 281 (Mammoth Cave Nat. Park, Kentucky).— PARKER,
Jour. Tennessee Acad. Sci., 12, no. 1, Jan., 1937, p. 85, fig. 18 (Bayou
du Chien; Reelfoot Lake, Tennessee); Rep. Reelfoot Lake Biol.
Sta., 3, Jan. 1939, p. 88 (Tennessee: Reelfoot Lake).— CAHN,
Illinois Biol. Monogr. 16, Aug. 31, 1937, p. 184, pis. 25-27, 30 fig. b,
map 20 (Illinois).— GREETE, Herpetologica, 1, no. 4, Nov. 16,

1937, p. 116 (West Virginia: Randolph Co., Tygart River near El-
kins). — ALEXANDRE, Soc. Canadienne Hist. Nat., tract no. 39,
Apr. 1937, p. 2 (Canada: Quebec: Lake Champlain and Richelieu
River, tributary to St. Lawrence R., right side).— HENNING,
Copeia, 1938, no. 2, June 30, 1938, p. 92 (Boone Co., Missouri).—
DENNING and BLACK, Occas. Pap. Univ. Arkansas Mus., No. 1,
June 1938, p. 46 (Arkansas: Lafayette, Prairie, Chicot, Clay, Gar-
land, Lawrence, and Washington Cos.). — CONANT, Amer. Mid-
land Natural., 20, no. 1, 1938, p. 157, pi. 21, fig. 1 (right), pi. 22,
figs. 1, 2 (Ohio: map); Herpetologica, 1, no. 5, Dec. 30, 1938, p. 138
(Ohio: Lucas Co.).— NECKER, Bull. Chicago Acad. Sci., 6, no. 1,
1939, p. 10 (Illinois: Cook Co.: Evanston; Kankakee Co.: Kankakee
Riv. near Altort).— WELTER and CARR, Copeia, 1939, no. 3,
Sept. 9, p. 130 (Kentucky: Triplet Co.; Fox, Fleming Co.; rare in
East Ky.).— LOGIER, Canad. Field-Nat., 39, May, 1925, p. 95
(Point Pelee; Hamilton, Dundas Marsh, Ontario, Canada); Roy.
Ontario Zool. Mus. Handb. no. 4, 1939, p. 56, pi. 7, fig. 2; pi. 8, fig. 7
(Ontario: Hamilton Bay and Dundas Marsh, Wentworth Co.;
Thames River at Beachville, Oxford Co. ; Grand River at Dunnville,
Haldimand Co.; Long Point, Norfolk Co.; and Point Pelee, Essex
Co.).— GENTRY, Rep. Reelfoot Lake Biol. Sta,, 5, Jan. 1941, p.

stejneger: American soft-shell turtles 49

75 (Tennessee: Clay Co., Overton Co.); Journ. Tennessee Acad.
Sci., 16, no. 3, 1941, p. 332 (Tennessee: Overton, Fentress, Pickett,
Jackson, and Clay Cos.).— CAGLE, Copeia, 1942, no. 3, Oct. 8,
p. 155 (Illinois: Jackson and Williamson Cos.)
Platypeltis spinifera RUTHVEN and THOMPSON, Herpet. Michi-
gan, 1912, p. 129 (Michigan).— THOMPSON, Misc. Pap. Zool.
Michigan, 1916, p. 63 (Monroe Co., Michigan).

1827.— Trionyx ocellatus LESUEUR, Mem. Mus. Hist. Nat. Paris, 15, Dec.
1827, p. 261 (type-locality, New Harmony, Indiana).— DeKAY,
Zool. New York, pt. 3, Reptiles, 1842, p. 7 (Mohawk River, New
York).

1838. — Trionyx annulifer WIED, Reise Nord-Amerika, 1, pt. 3, p. 140 (type-
locality, Ohio River at Pittsburgh, Pennsylvania); Voy. Amer.
Nord., 3, 1843, pp. 242-243.

1844.— Tyrse argus GRAY, Cat. Tort. Brit. Mus., p. 48 (type-locality, "West
Africa, Sierra Leone?"; type in Brit. Mus.; Lord Derby, collector);
Knowlsley Menag., 1846 (pi. — ).
Trionyx argus GRAY, Cat. Shield Rept. Brit. Mus., pt. 1, March 8,
1856, p. 68.

1856. Trionyx annularis "Wied", GRAY, Cat. Shield Rept. Brit. Mus., pt.

1, March 8, 1856, p. 69 (lapsus in synonymy).

1857. Aspidonectcs nuchalis AGASSIZ, Contr. Nat. Hist. United States, 1,

p. 406 (type-localities, Cumberland and Tennessee rivers; cotypes,
Mus. Comp. Zool., nos. 1623-1625, Cumberland river).— COPE,
Bull. U. S. Nat. Mus., no. 1, 1871, p. 51.— TRUE, Bull. U. S. Nat.
Mus., no. 24, 1883, p. 5; Fish. Industr. United States, sect. 1, 1884,
p. 152 (middle western states).— DAVIS and RICE, Illinois State
Lab. Nat. Hist., Bull. No. 5, 1883, p. 52 (Cumberland and Upper
Tennessee Rivers).— JORDAN, Man. Vert. Anim. North United
States, ed. 5, 1888, p. 206 (Cumberland and Upper Tennessee Rivers) ;
ed. 8, 1899, p. 206.— RHOADS, Proc. Acad. Nat. Sci. Philadelphia,
1895, p. 404 (Cumberland River, near Nashville).

1861. Amyda mutica MILES, First Biennial Rep. Geol. Surv. Michigan, pp.
2323, 233 (Michigan). (Not of Lesueur).— SMITH, Science News
Suppl., 1, 1879, p. 7 (Michigan).— KIRSCH, Bull. U. S. Fish Comm.
1894 (1895), p. 41 (Indiana: Eel River); p. 333 (Maumee Riv.,
Ohio).

1865. — Gymnopus olivaceus WIED, Nova Acta Acad. Leopold.-Carol., 32,
pt. 1, p. 55, pi. 5 (type-locality, New Harmony, Wabash River,
Illinois).

1869.— Callinia spicifera GRAY, Proc. Zool. Soc. London, 1869, pp. 222
(lapsus for spinifera).

50 bulletin: museum of comparative zoology

1939.— Amyda spinifera spinifera STEJNEGER and BARBOUR, Check
List N. A. Amph. Rept., ed. 4, p. 172; ed. 5, 1943, p. 213.— ANDER-
SON, Bull. Chicago Acad. Sci., 6, no. 11, July 8, 1942, p. 219 (Mis-
souri: Jackson Co.; Missouri River near Atherton). — EDGREN,
Copeia, 1942, no. 3, Oct. 8, p. 180 (Michigan: Van Buren Co., Rey-
nold Lake).— PETERS, Copeia, 1942, no. 3, Oct. 8, p. 183 (Illinois:
Cumberland Co.).

Geographical distribution

Amyda spinifera has been credited to the fauna of South Carolina
on the strength of the specimen U.S.N.M. No. 7650 recorded in U. S.
Nat. Mus. Bulletin No. 24 (1883), p. 29, as coming from "Abbeville,
S. C". It is also the record upon which is based True's record of the
occurences of Aspidonectes spinifer in H. S. Thompson's, South Caro-
lina, 1883, p. 238. The record is undoubtedly erroneous, the ascertain-
able facts being as follows :

The specimen, recently hatched, was one of the numerous neglected
turtles found in the collection when Dr. G. Brown Goode took charge
and tintagged and registered the specimens. He entered the present
one in 1872 as No. 7650, the tin tag bearing that number. In the record
book [in Brown Goode's handwriting] it is noted as an Aspidonectes
[no specific name] ; locality : "Abbeville S. C"; and no further remarks.
There is now no other indication of its origin than an old torn scrap
of paper in the bottle with "7650 Abbeville S. C." likewise in his hand-
writing. On the back of this label, however, there is — in very faded
ink and in an entirely different handwriting — the remnant of an in-
scription, beginning and end clipped off, "ville Mis". If this remnant
really is part of the original locality record, it seems probable that it
may have read "Abbeville, Miss." and inadvertently attributed to
South Carolina as the more familiar locality. The specimen is unques-
tionably an Amyda spinifera. The whole pattern of coloration is
normal of that species, which of course at once excludes A. ferox. The
single regularly defined submarginal dusky ring on the upper side of
the disc likewise excludes A. agassizii, the species one would expect
if it were collected in Abbeville, S. C. To make perfectly sure of its
identity I have had the skull extracted. It shows all the characteristic
features of A. spinifera as distinguished from A. agassizii.

Another erroneous record of Trionyx spiniferus, viz. in Pope's
Turtles (1931) pi. 45, figs. 98, 99) showing excellent figures of two soft-
shelled turtles from the region of Columbia, South Carolina, needs
correction. Both pictures are plainly of Amyda agassizii.

stejneger: American soft-shell turtles

•

art

V|_

t/3

■*-'

Ill

c
O

Average

of 9
Specimens

co
m

101386

Virginia:

Smyth Co.;

Seven Mile

Ford

59263

Minnesota

Winona

Co.;
Homer

59264

Minnesota

Winona

Co.;
Homer

o
m

54730

Iowa:

Muscatine

Co.;

Fairport

p
in

54731

Iowa:

Muscatine

Co.;
Fairport

o
m

59265

Minnesota
Winona

Co.;
Homer

54421

Montana:

Big Hern

Co.;

(Vow

Agency

o
©

70397
Oklahoma:
McCurtain

Co.

54732

Iowa:

Muscatine

Co.;
Fairport

O
CO

Basicranial length .... mm.

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52 bulletin: museum of comparative zoology

Stockwell's reference to the occurrence of Aspidonectes spinifer
"North of Athabasca Lake" (Journ. Compar. Medic. Surg., 9, 1888,
p. 28) must rest on some curious lapsus.

Where Amy da spinifer meets with A. asper is as yet conjectural.
It extends at least as far south as the northern part of the State of
Mississippi, for we have undoubted specimens from De Soto County
(U.S.N.M. No. 92606) and Lake Washington, Washington County
(No. 92607). A young male specimen from Madison Co., Northern
Louisiana (No. 83985) is likewise this form with only one marginal
stripe.

The Museum of Comparative Zoology has a specimen from Colum-
bus, Ga (M.C.Z. 1606).

Mississippi River and tributaries, west to Colorado, north to Mon-
tana; St. Lawrence River and tributaries; east to Vermont, western
New York, and Pennsylvania.

A. nuchalis, judging from the slight material at hand, is not a
strongly differentiated form. The character chiefly relied on by Agas-
siz, "the most prominent specific character . . . the marked depression
on either side of the blunt median keel," does not hold in a series of
specimens. I find quite a number so characterized among typical A.
spinifer a, and while one of the specimens from near Sevier ville, Tenn.,
has a rather flat bony carapace, the other has "the blunt keel, which
extends along the median line and slopes uniformly upon the sides,"
exactly as he describes it diagnostically for A. spinifera (p. 404).
The angle of the entoplastron, in the few examples examined, is some-
what more obtuse, between 90° and 95°, than in corresponding speci-
mens of typical A. spinifera. The sharply defined ocelli on the carapace
seem to be larger than in A. spinifera of the same age (size).

I do not at all understand Agassiz's note that "this species differs
strikingly from Asp. spinifer in the much more elongated form of the
male, and in the great development of the marginal spines and of the
tubercles upon the carapace, which project very slightly in the male
Asp. spinifer." On the contrary, in our upper Tennessee specimens,
presumably typical A. nuchalis, the carapace of the males is wider
than in the females, and the spines on the anterior edge very much
smaller, exactly as in typical A. spinifera.

U.S.N.M. 86677 from Cumberland Gap, and 86682 from 2 miles
west of Sevierville, Tennesse, are typical "nuchales."

stejneger: American soft-shell turtles

List of specimens in the U. S. National Museum

Mont., Ft. Union

F. V. Hayden

7163(029528)

Tenn., Nashville

J. Varden

7165(029529)

it it

7166(029530)

it a

7167

n it

7169 juv.

tt tt

7648 & adol.

Wyo., Ft. Laramie

F. V. Hayden

7649

7650 pull.

Miss., Abbeville?

7661 juv.

111.

R. Kennicott

8359

Ind., Madison ?

9654

111., Mt. Carmel

R. Ridgway

May,

1878

9717

111., Mt. Carmel

Mrs. L. M. Turner

June,

1878

9928 juv.

Iowa, Webster City

C. Aldrich

1878

11625 juv.

11631 juv.

La., Prairie Mer Rouge

12061 juv.

111., Mt. Carmel

L. M. Turner

14535 juv.

Mont., Ft. Custer

C. Bendire

14536 9 ad.

it a

16704 9

Ala., Courtland

rP. H. Kirch, E. O.

■

Jones, and

May,

1889

16705

ti a

W. M. Andrews

17823 juv.

Ark., Benton

Jordan & Gilbert

19622-3 9 & adol.

19625

21128-9 juv.

Ohio, Cuyahoga River

A. J. Woolman

July

25, 1893

21416-7

21567-8

Ohio, Edgerton

P. H. Kirsch

July

28, 1893

21569-70

Ind., Fish Cr., near

Hamilton

July

21, 1893

21571-7

Ohio, Maumee Basin

1893

22711

Ind., Vincennes

R. Ridgway

24536

Mont., Ft. Custer

C. Bendire

Mar.

8, 1886

26290

Ohio, Franklin Co.

R. C. Osborn & E.

C. Williamson

June,

1897

029014 9

" Columbus

O. Davie

33494

Ind., Lake Maxin-

kuckee

B. W. Evermann

July

21, 1900

33495

Yellow River nor(

of Burr Oak

Oct.

3, 1900

33496-501

" L. Maxinkuckee

1899-

-1900

33767

W. Va., Dry Fork,

Perryville

W. P. Hay

1900

34 bulletin: museum of comparative zoology

35404-8

Ind., L. Maxinkuckee

H. W. Clark

1900

36412

111., Illinois R.

S. P. Bartlett

42583 juv.

Ind., Long Point, L.

Maxinkuckee

Evermann & Clark

Oct.

5, 1906

42584 juv.

" L. Maxinkuckee

n a

42585 eggs

u a

a it

Nov.

17, 1906

42905-6 juv.

" Burlington

B. W. Evermann

1899

50670 juv.

Twin Lakes

H. W. Clark

July

2, 1909

51213 hf-gr.

Mich., Monroe Piers

C. Rutter

Aug.

13, 1894

51214 "

Ohio, Toledo, Grassy

Point

Aug.

3, 1894

51529 ad.

Kansas

R. L. Moodie

53521

Iowa, Fairport

Bur. Fisheries

Apr.

24, 1911

53522

111., Hamilton

J. McAdams

May

1, 1915

53523-6

Iowa, Fairport

E. Snyder

Oct.

1, 1914

54421 cf ad.

Mont., Crow Agency

M. A. Hanna

Aug.

5, 1916

54422-3

it a

R. Kellogg

July

8, 1916

54730 9 adol.

Iowa, Fairport

J. Snyder

June,

1916

54731 9 adol.

a tt

July

2, 1916

54732

a a

54739-41

n a

May

8, 1916

54743-6

a a

May,

1916

54747

111., Meredosia

Freeland & Williams 1908

55680

" Madison Co.

J. Hurter

55681

" Union Co.

55682

Ky., Morgan Co.

'55683

Kans., Greenwood Co.

July-Aug., 1912

55684

Mo. Stone Co.

June 27, 1908

55085

" St. Louis Co.

1913

55686

" St. Louis

55687

it a

Apr.

9, 1905

55688

" Reynolds Co.

July

21, 1911

55689

" FranklynCo.

Aug.

20, 1911

55690

" Washington Co.

June

25, 1911

59046

" St. Louis

May

20, 1908

59263-6 9 adol

. Miss., Homer

F. Schrader

59267 9

Mo., Alexandria

E. Stringham

June

5, 1916

59270-5

Minn., Homer

F. Schrader

Sept.

8, 1916

59277

111., between Warsaw

and Hamilton

E. Stringham

Aug.

25, 1916

59279-80

Mo., Canton

Bur. Fisheries

59285

Iowa ?

59736

Mont., Crow Agency

R. Kellogg

July

23, 1916

59956

Ind., Madison

O. P. Hay

59979 d" juv.

Ind. ?

stejneger: American soft-shell turtles

60571 juv.

III., Madison Co.

J. Hurter

70397 adol.

Okla., Red River,

McCurtain Co.

A. I. Ortenburger

1924

72387

Ind., Knox

H. W. Clark

Aug.

12, 1909

73668-9 juv.

Miss., Greenwood

I. L. Towers

1925

83985 <? adol.

La., 2 mi. E. of Mounds C. E. Burt

June

30, 1931

86677 9 ad.

Tenn., 5 mi. S.E.

Cumberland Gap

July

22, 1932

86681-2 d" adol

.. " 2 mi. W. Sevierville

1932

87164 juv.

" 2 mi. S. Kingston "

July

27, 1932

90441-4 juv.

Kans., 1 mi. W. Winfield

Apr.

30, 1933

91022 juv.

" Winfield

L. Hoyle

May

28, 1933

92606 pull.

Miss., Lake Cormorant S. F. Hildebrand

June

9, 1933

92607 juv.

" Greenville

May

31, 1933

93089-94 juv.

Mo., Dardenne Cr.,

St. Charles Co.

L. Hubricht

Aug.

16, 1931

95140 pull.

Miss., 1 mi. W. Yazoo

City

C. E. Young

Aug.

13, 1934

95261 9 ad.

Kans., 2 mi. E. of

Calista

C. E. Burt

May

25, 1934

95301

" 11 mi. S.E. of

Winfield

Aug.

31, 1934

95352 9 ad.

Ark., 7 mi. N.W.

Natural Dam

June

7, 1934

95405 & ad.

Mo., Glaize Creek,

Jefferson Co.

A. A. Heinze

July

20, 1932

99862-75

La., Red River near

Shaw

C. E. Burt

1935

100160 9 ad.

" Bayou Chene

June

17, 1935

100202-12

" False River near

New Roads

June

17, 1935

100420-1

" Cane River near

Natchitoches

June,

1935

100529-30 & 9

Kans., Winfield

Aug.

12, 1935

100580

" Lake City

June,

1935

100795 pull.

Ind., Irvington

O. P. Hay

101386 9 ad.

Va., Seven Mile Ford

A. Wetmore

June

1, 1935

102705 juv.

Miss., Belzoni

S. F. Hildebrand

July

10, 1936

102911 c? juv.

Tenn., Reelfoot Lake

W. M. Perrygo &

C. Lingebach

May

8, 1937

102912

n u u

W. M. Perrygo &
C. Lingebach

103477-9 juv.

Vt., Swanton

L. H. Babbitt

June

21, 1937

bulletin: museum of comparative zoology

107786

107787

109178 pull.
113228 juv.
115980 pull.

Term., Iron Creek near

Willow Grove G. Gentry

mouth of Wolf

River
La., Jonesville G. K. Payne

July 17, 1939

July 19, 1939
June, 1940

Miss., Deer Creek C. Hollingsworth June 24, 1940

"Amyda spinifera aspera1 (Agassiz)
Plates 17, 18, 19c

1854.— Trionyx ferox WAILES, Rep. Agric. Geol. Mississippi, pp. 327, 331

(Mississippi) (not of Schneider). — Plalypeltis ferox AGASSIZ, Contr.

Nat. Hist. United States, 2, 1857, pi. 6, fig. 3 (Mobile, Ala.; Mus.

Comp. Zool., no. 1608A).— Aspidonectes ferox JORDAN, Man.

Vert. North Amer., ed. 5, 1888, p. 206 (part).
1857. — Aspidonectes asper AGASSIZ, Contr. Nat. Hist. United States, 1, p.

405 (type locality, Lake Concordia, La.; cotypes, U.S.N.M. no.

012349, Prof. B. L. C. Wailes, collector, and Mus. Comp. Zool, no.

37173).— COPE, Bull. U. S. Nat. Mus., no. 1, 1875, p. 51 (Lower

Mississippi tributaries). — DAVIS and RICE, Illinois State Lab.

Nat. Hist., Bull. no. 5, 1883, p. 52.— TRUE, Fish. Industr. United

States, sect. 1, 1885, p. 152.— BAUR, Amer. Natural., 22, 1888, p.

1122; Proc. Amer. Philos. Soc, 31, July 1893, p. 217.— BEYER,

Proc. Louisiana Soc. Nat. Hist., 1897-1899 (1900), p. 43 (Louisiana).
1892. — Trionyx agassizii HAY, Indiana Geol. 17 Rep., p. 552 (part); Batr.

Rept. Indiana, 1893, p. 144 (part) (not of Bauer 1888).
1899. — Aspidonectes agassizi JORDAN, Man. Vert. North Amer., ed. 8, p. 206

(part only) (emendation) (not of Bauer).

1919. — Amyda spinifera BABCOCK, Mem. Boston Soc. Nat. Hist., 8, no. 3,
pi. 32 (not of text; not of Lesueur).— HALTOM, Alabama Mus. Nat.
Hist., Mus. Pap. no. 11, 1931, p. 142 (Alabama: Marengo Co.: near
Demopolis, Tombigbee River).

1923.— Trionyx spiniferus agassizii SIEBENROCK, Verh. Zool. Bot. Ges.
Wien, 72, Aug. 1923, p. 188 (part: West Louisiana).

1939.— Amyda spinifera aspera STEJNEGER and BARBOUR, Checklist N.
Amer. Amph. Rept., ed. 4, p. 172 (Lower Mississippi tributaries in
Louisiana and Mississippi) ; ed. 5, 1943, p. 213 (Lower Mississippi tri-
butaries in Louisiana; rivers of Mississippi and Alabama).

1 Latin: rough; probably with reference to the "prominent warts of the bony plates," which,
according to Agassiz, "exist in no other species with which I am acquainted."

stejneger: American soft-shell turtles 57

•

Types and type locality. The two specimens specifically mentioned
by Agassiz as basis for his A. as per were "an imperfect skeleton . . .
belonging to the Smithsonian Institution and prepared from a speci-
men forwarded by Professor B. L. C. Wailes of Washington, Missis-
sippi," and "a stuffed specimen belonging to the Museum of the
University of Oxford, that has been collected during the Geological
Survey of Mississippi, under the superintendence of Professor
Wailes."

In a letter dated Washington, Miss., 8: January, 1853, Professor
B. L. C. Waile wrote to Professor S. F. Baird that he had forwarded
to him a number of reptile specimens, among them "2 shells and
crania of Trionyx ferox." They were entered in the register of the
osteological collections of the Smithsonian Institution under the
generic name Trionyx only by Professor Baird on March 21, 1853 as
numbers 1084 and 1086, received from' 'B. L. C. Wailes, Washington,
Miss." Of these specimens there are now in the National Museum: 1)
a skull marked 1084; 2) a skull, with carapace and plastron in pieces
(entoplastron and epiplastra missing), marked 108G; and 3) a carapace
numbered 12349. The specimens have no original labels attached to
them, but the numbers on the skulls are written on them in ink, and
on the inside of the carapace of No. 12349 there is written with black
ink in the same "professional" hand characteristic of all the specimens
received from Wailes: "Trionyx ferox? Lake Concordia, Louisiana,
1851, B. L. C. Wailes." This is undoubtedly the "imperfect skeleton"
examined by Agassiz; it is characterized by the "prominent warts of
the bony plates" on the posterior part of the bony disk described by
Agassiz (I.e. p. 406), which "bony warts exist in no other species with
which I am acquainted."1

The second cotype, the "stuffed specimen" received from the Uni-
versity of Oxford, Mississippi, is in the Museum of Comparative Zoo-
logy (M.C.Z. no. 37173) where I was permitted to examine it. It is an
adult male, inscribed in the same bold, handsome style as Wailes'
other specimens Trionyx ferox, but no locality data. I made the fol-
lowing notes: "Bony disk 200 mm. long, 185 mm. wide; 7 neurals,
seventh small; sculpture fine, of the spinifer style, with the bony
tubercles on the seventh pleural described by Agassiz, tubercles on the
posterior and anterior flaps of leathery disk, spinous tubercles on edge
of carapace anteriorly between legs; callosities on plastron large, al-
most meeting on the mid-line; a median large triangular callosity

1 Agassiz did not know the bony disk of the true A. ferox from Florida.

58 bulletin: museum of comparative zoology

(sides about 25 mm.) on the entoplastron ; entoplastral angle slightly
less than 90°.

The specimens which Agassi z mentions as having been received
"through the kindness of Mr. Winthrop Sargent of Natchez"1 may be
regarded as paratypes. One of them, a very large male (M.C.Z. no.
1597), I have examined and made the following measurements and
notes :

Total length of leathery carapace 450 mm.

Width of leathery carapace 370

Length of bony carapace 240

Width of bony carapace (at pleural 4) 240
Height of body (at neural 1) 88

Large rounded tubercles on front edge of leathery carapace; flat
tubercles of about same size on front and hind flaps; callosities on
xiphi-plastra meeting; closest approach of hyo-hypoplastral callosities
8 mm. ; bony ridges comparable to those on bony carapace of U.S.N.M.
no. 1086; sculpture somewhat coarser (because of larger size of speci-
men) ; entoplastral angle about 90°. Distinct traces of two black mar-
ginal rings on hind flap of carapace.

A young paratype, probably of the same origin (M.C.Z. no. 1622)
with a recent label in lead pencil "No. 1622. Type Amyda asper
(Agassiz) Lake St. John, Miss.2 W. Sargent leg. et don." measures ap-
proximately 65 mm. in length and 57 mm. in width. The secondary
marginal lines on the posterior flap of the carapace characteristic of
normal individuals of the subspecies of corresponding age are repre-
sented by two series of closely set spots (pi. 16, fig 1).

Notes on synonymy. Special attention is called to the reference to
Agassiz's illustration (Contrib., 2, pi. 6, fig. 3) of a young specimen of
A. aspera under the name PlatypeHis fcrox, as it apparently has given
rise to great confusion among southern herpetologists who, because of
it, have mistakenly identified young specimens from Alabama, Missis-
sippi, and Louisiana. The young of the true A. ferox was unknown to
Agassiz, but on account of the locality (Mobile) of the specimen figured
and its similarity in color pattern to certain soft-shell turtles from
South Carolina he assumed that it represented the Florida species.
The original of the figure 3 appears to be still at the Museum of Com-
parative Zoology bearing the number 1608A. It turns out to be a

'Agassiz, in 1856, had J. Burkhart make colored illustrations from a female of the same lot
of specimens which shows plainly the marginal black rings on the posterior flap. Reduced
copies of the drawings are herewith presented (pis. 20, 21) through the kind permission of
Dr. Thomas Barbour, Director of the Museum of Comparative Zoology.

2 Possibly a slip for Louisiana.

stejneger: American soft -shell turtles 59

young of Agassiz's own Aspidonectes as per, and the figure is herewith
reproduced under its proper name (pi. 19, fig. c) for comparison with
the young of the true A. ferox (pi. 19, fig. a and b).

Babcock's figure of A. spinifera, as quoted above, does not represent
the true nominate form of that species, but is either an A. spinifera
aspera or an A. agassizii. Dr. Babcock, in a letter of December 13,
1933, kindly informs me that the picture was made, during his absence
in 1918, from a living specimen, the origin and disposition of which
he has been unable to trace. The exact identification depends some-
what on the artist's accuracy. My best guess is that the specimen was
an A. aspera, judging by the extreme length of the proboscis.

Variation. A series of four living young adults were presented in
September, 1934, by Mr. S. E. Brand, of Canton, Mississippi, all taken
in barrow pits, three on Pearl River and one on Big Black River. The
specimens unquestionably represent the same form and, as highly
instructive, some of the notes made at the time may be of interest, the
color designations in quotation marks having reference to Ridgway's
Nomenclature of Colors, 1886: —

U.S.N.M. no. 95191, 9 ad. Iris clear "primrose yellow" with black
horizontal bar not quite reaching pupil. General color above "tawny-
olive," head, neck, and legs densely speckled with dark brown, cara-
pace with obscure, irregular blotches of "raw umber," in the center
of which one or more small blackish spots form interrupted ocelli, the
two outer rows of which posteriorly assume the form of short lines
parallel with the submarginal ring; marginal edge dull "olive-buff,"
top of head like carapace with the shallow fork on the snout chiefly
indicated by the black outlines; side of head like top down to the lips;
a slightly paler postocular band tinged with yellow and narrowly but
very distinctly margined with black, the upper line continuing back-
ward the whole length of the neck ; upper side of legs slightly paler than
carapace and more tinged with olive; under side pale flesh color caused
by the fine network of blood vessels shining through the white ground
color; plastral callosities pale "vinaceous-cinnamon" more or less
tinged with bluish; under side of feet tinged with "verditer blue" with
a wash of yellow deepening towards the outer margin, inner half
spotless, outer half, including web, marked with heavy blackish anas-
tomozing lines and spots; claws yellowish white; under side of neck
faintly mottled with obscure "pinkish vinaceous" spots; throat
strongly washed with "pale blue"; lips and under side of proboscis
tinged with "gallstone-yellow." Tubercles on anterior edge of cara-
pace triangular, pointed, about 3 mm. long; anterior flap densely

GO bulletin: museum of comparative zoology

crowded with smaller and blunter tubercles of various sizes; posterior
flap with larger, oblong, blunt tubercles, fairly regularly spaced; cara-
pace posteriorly with similar but lower and longer tubercles in fairly
regular longitudinal series. Skin covering nuchal of bony carapace
with a median longitudinal series of about 4 enlarged tubercles ex-
tending on to anterior leathery flap. Fontanelles between nuchal and
first pair of pleurals covered with smooth skin upon which are a few
tubercles. Callosities covering plastral bones rather small, leaving
median fontanelles larger than xiphiplastral callosities; no trace of
callosities on entoplastron and ectoplastron ; entoplastral angle 90°.

U.S.N.M. no. 95192, also from a "barrow pit" at the nearby Big
Black River, a young adult 9 , smaller than 95191, is structurally al-
most identical with it, except for the smaller dimension and being
slightly broader. The coloration of the living specimen was darker and
more pronounced. Iris "ochraceous buff," black horizontal bar reach-
ing pupil. Carapace "raw umber" with large irregularly buffy-edged
blotches of "mummy brown," increasing towards and infringing upon
the marginal "buffy" edge and its bordering dusky ring; towards the
margin the blotches coalesce more or less so as to form three distinct
but interrupted submarginal rings; color of upper side of neck, head,
and «legs like that of the carapace but more "tawny" and thickly
sprinkled with heavy blackish spots ; forked figure on top of snout deep
"ochraceous buff," with very distinct black edges like the ochraceous
postocular stripe; under side milk white strongly suffused with flesh-
color; palms and soles strongly tinged with tawny, palms heavily
marked with coarse blackish anastomozing lines and spots, soles al-
most devoid of markings, claws white, throat tinged with pale blue;
callosities "vinaceous-cinnamon" with slight tinge of blue in center.

U.S.N.M. no. 95193, young adult d\ from Pearl River, differs
structurally from the two described females in showing no trace of
fontanelles on carapace, in much larger plastral callosities than on
xiphiplastra nearly meeting in the middle, those covering hyoplastra
and hypoplastra only 4 mm. apart; in addition there is a large triangu-
lar callosity on the entoplastron ; the row of tubercles on the edge of
anterior flap of carapace barely indicated and no tubercles on flap and
nuchal. In addition, the skin of the carapace, bony disk as well as
lateral and posterior flaps, is densely sprinkled with very minute hard
tubercles which make the skin feel like fine sandpaper. The color
notes on the living specimens are as follows : Iris pale buff, black bar
scarcely separated from pupil. Upper side nearly uniform "tawny
olive," the region of the bony disk distinctly more olive; small, round,

stejneger: American soft-shell turtles 61

dark brown spots of varying size scattered sparingly over the carapace;
the normal margin not perceptibly lighter than rest of carapace and
barely set off from it by the obscure dusky submarginal ring; no in-
dications of additional rings by dark lines or spots; top of head and
neck like back with minute black scattered dots; lips and sides of neck
strongly washed with "gallstone yellow," with normal postocular
stripe pattern indicated; fork figure on snout normal, buff colored,
black-edged; legs above and feet coarsely spotted with black; upper
side of tail like feet with lateral blackish lines converging backwards.

U.S.N.M. no. 95194, from the same locality, is also a male, and
slightly smaller. It shares the same structural characteristics, only
the tubercles on the anterior edge of carapace are slightly larger and
more distinct; the outline of the soft carapace is more oval than
rounded ovate; the xiphiplastral callosities are continuous, but the
distance between the hyohypoplastral callosities is a trifle wider, and
the entoplastral callosity somewhat smaller; the "sandpaper" effect
of the back is very much alike, but a double series of larger flat tuber-
cles on the mid-line of the 7th pleural is quite noticeable. The colora-
tion is also much the same, only the ground color of the carapace is
lighter, and the dark spots distinct, the chief difference being the inter-
rupted rows of the three marginal rings; on the other hand, the dusky
pattern elsewhere is finer; the pale edge of the leathery disk is slightly
paler than the rest of the disk.

For an easier appreciation of the more striking variations, they are
summarized in table 7. The differences are of various significance. As
certainly indication of sex is the greater development of the plastral
callosities of the males, besides the well-known difference in the
length of the tail, the "sandpaper" effect seems to be peculiar to a cer-
tain age (or size, or season?) of young males; the greater development
of the tubercles or "spines" may be correlated with sex or with age or
with both, remembering that the females are larger than the males;
the closing of the nuchal fontanelles of the carapace seems to take
place much earlier in the males than in the females; the slight differ-
ence in the outline of the leathery disk (as seen from above) is not due
to sex, and, at the stage of the specimen under discussion, not to age,
though the general rule in these turtles is that the young are more
circular than the old ones. None of the above characters seem to be
of specific or subspecific significance. On the other hand, the presence
of two or more concentric blackish rings inside the normal Amydan
dusky ring delimiting the pale rim of the leathery carapace is of diag-
nostic value in defining the subspecies, but unfortunately it has a

bulletin: museum of comparative zoology

tendency to be obscured or absorbed with age. Individual specimens
may occasionally be found without these additional rings among nor-
mal populations, but they are the "intermediates" which justify the
use of the trinominal nomenclature. On the other hand, it is significant
that among a very large number of specimens from northernmost
Montana to Louisiana and from Colorado to Lake Champlain, I have
not seen an authentic specimen of Amy da spinifer with two or more
submarginal rings.

Geographical distribution

Lower Mississippi tributaries in Louisiana and Mississippi and
Alabama.

Table 7
Variations in young adult A. aspera from Mississippi

Mississippi

Pearl

Black

Pearl

River

95191

95192

95193

95194

Sex

Soft carapace length mm.

286

210

180

158

width mm.

242

182

152

140

Bony carapace length mm.

173

119

111

width mm.

131

110

Plastral callosities

Small

Large

Callosity on entoplastron

Spines on edge of carapace

Large

Median

Small

pointed

blunt

Tubercles on median line of front carapace

flap

Carapace skin "sandpapered"

Carapace fontanelles

Outline of carapace (from above) oval

" " " " " ovate

stejneger: American soft-shell turtles

Intergrades aspera-spinifera

It has been repeatedly asserted that the only differences by which
specimens of A. asper a can be recognized in the adult state are "the
very coarse and large tubercles of the front and hind part of the cara-
pace, which extend, behind, even over the bony shield, and are there
supported by prominent warts of the bony plates" (Agassiz, Contr.,
1, p. 406) and by the fact "that in younger specimens of Asp. asper
there are . . . two or three black lines separating the pale rim of the
posterior margin, whilst there is only one in Asp. spinifer." (Agassiz,
I.e.). As Agassiz himself observes, these lines fade away "pretty soon."

Table 8
Cranial measurements of A. asper a in millimeters

Basicranial length

mm.

Tip of snout to orbit
Horizontal diameter of orbit
Orbit of tympanic cavity
Longest diameter of temporal fossa
Interorbital width
Width of maxillary alveole
Length of internal choanae
Internal choanae to intermaxillary

foramen
Length of intermaxillary foramen
Length of mandibular symphysis
Width of mandibular alveole surface

C
o

CO
00

63 63

16
12
19
11

5.5
10

16
10.5
18.5
12

6.5
6.2

o
>-

C
o3

O
i— i

16
12
19
13

5.5

6.5

"oj

1-5

00
CD

10
8
9

6.5
3
3
6

6
5
6
2.5

23 69

5.5

3
3
3
2

o3
C
c3

3
O

h-1

CO
CO
<M
Oi
CM

19
13
20.5
13

6
10

6.5
7
11.5
5.5

02
CO

bD
cj
s-,

53.0

13.8
10.1
15.3
9.9
4.4
4.4
8.1

5.9
5.5
6.4
3.5

bulletin: museum of comparative zoology

Specimens of intermediate ages are difficult to identify with our present
knowledge. Apparently the double or triple rings are not always a
constant character. I have before me, through the courtesy of Miss
Fannye A. Cook, a specimen collected about a mile or two southeast
of Brookhaven, Lincoln Co., Mississippi, in a small tributary of the
Bogue Chitto, Pearl River drainage. Its carapace measures only 43
mm. in length, hence it is quite young. It seems to be an aberrant
specimen of asper a, the anomaly being in the absence of the second
dark ring on the carapace margin. From its locality it ought to be
aspera, but such abnormal (or "incompleted" or "reversed") specimens
are known.

List of specimens in the U. S. National Museum

01084

Miss., Washington?

B. L. C. Wailes

01086

H 11

Cotype of
Aspidenectes asper
Agassiz

7653-4

" Monticello

H. Tennison

012349

La., Lake Concordia

B. L. C. Wailes

1851 Cotype of
Aspidonectes asper
Agassiz

13250 9

" New Orleans

R. W. Shufeldt

1883

029266

S. W. Harvey

029310 ad. 9

" near New Orleans

U. S. Fish Comm.

66147 juv.

" Madisonville

May 29, 1886

68054 cf adol.

" Roberts

R. F. Shaw

79350-1 juv.

Miss., 1 mi. W. of Mel-

R. Kellogg and

vin

N. Boss

Oct. 1929

83996 9 juv.

Ala., 3 mi. S.E. of

Coatopa

C. E. Burt

July 1, 1931

95191 9 ad.

Miss., Pearl River

S. E. Brand

Aug. 1934

95192 9 adol.

" Big Black River

95193 d" adol.

" Pearl River

95194 d" adol.

u ii

100650 c? ad.

La., near Atchafalaya

C. E. Burt

June 17, 1935

100805 pull.

Miss., Enterprise

O. P. Hay

1881

115979 9 adol.

" near Guntoun

E. & W. H. Patten

Aug. 4, 1940

115981 juv.

" Chookatonkchia

Creek

H. L. Owens

June 30, 1941

stejneger: American soft-shell turtles 65

Amyda emoryi1 (Agassiz)
Plates 24-25

1849. Trionyx ferox ROEMER, Texas, p. 171, p. 459 (at New Braunfels,
Texas, in the Guadalupe and Comal Rivers).

1857. — Aspidonectes emoryi AGASSIZ, Contr. Nat. Hist. United States 1, p.
407; 2, pi. 6, figs. 4-5 (type-locality, Rio Grande River, near Browns-
ville, Texas; cotypes, U. S. Nat. Mus. No. 7855; Mus. Comp. Zool.,
Nos. 1909, 1913; Dr. Kennedy, collector; Williamson Co., Texas). —
COPE, Bull. U. S. Nat. Mus., No. 1, 1875, p. 51 (Texas); No. 17,
1880, p. 13 (Dallas; Helotes Creek, near San Antonio, Texas). —
TRUE, Bull. U. S. Nat. Mus., No. 24, 1883, p. 29 (Matamoras,
Mexico; Texas: Brownsville, Rio Grande, Rio Seco, Braunfels; Old
Fort Cobb, Oklahoma); Fish. Industr. United States, sect. 1, 1884,
p. 152.— BEYER, Proc. Louisiana Soc. Nat., 1897-1898, p. 43
(Louisiana).— STRECKER, Trans. Texas Acad. Sci., 4, pt. 2, no.
5, p. 6 (McLennan Co., Texas); Proc. Biol. Soc. Washington, 21,
March 21, 1908, p. 79 (Bazos, Bosque River, McLennan Co., Texas,
abundant); 23, 1910, p. 121 (Bullhide Creek, McLennan Co., Texas).
Trionyx emoryi BOULENGER, Cat. Chel. Brit. Mus., 1889, p. 258 —
STRAUCH, Mem. Acad. Sci. St. Petersbourg, ser. 7, 38, no. 2, 1890,
p. 117 (Texas).— DITMARS, Reptile Book, 1907, p. 78 (tributaries
of the Rio Grande in Texas and Mexico).— SIEBENROCK, Zool.
Jahrb., Suppl. 10, pt. 3, 1909, p. 603 (Colmisneil, Tyler Co., South
Bosque Riv., Texas); Verh. Zool. Bot. Ges. Wien, 73, 1923, p. 190.—
LINSDALE and GRESSITT, Copeia, 1937, no. 4, Dec. 31, pp. 222-
225, figs. 1-3 (Colorado River: Delta, Lower California; Clark Co.,
Nevada; Mohave Co., Arizona; California Lakes, Imperial Co.,
California; transpl. to Colorado Riv.?). — LINSDALE, Proc. Ameri-
can Acad. Arts Sci., 73, no. 8, May 1940, p. 255 (Nevada: Clark Co. :
Colorado River).
Amyda emoryi STEJNEGER and BARBOUR, Checklist North Amer.
Amph. Rept., ed. 1, 1917, p. 124 (Rivers of Texas, north into south-
ern Oklahoma and Arkansas); ed. 2, 1923, p. 140; ed. 3, 1933, p. 153.
-PRATT, Man. Vert. United States, 1923, p. 249.— ? HAY, Pan-
Amer. Geol., 39, March 1923, p. 119, pi. 9, figs. 2-4 (fossil, Brazos
River at Pittbridge, Texas).— SCHMIDT, Copeia, no. 131, June 30,
1924, p. 64 (Arizona; introduced).— STRECKER, Baylor Univ.
Bull., 27, no. 3, Sept. 1924, p. 47 (eastern Oklahoma); Contr. Baylor
Univ. Mus., no. 2, Jan. 15, 1926, p. 3 (Somervell Co., Texas); no. 3,
Feb. 15, 1926, p. 4 (Liberty Co., Texas); no. 6, June 15, 1926, p. 8

1 To Col. Wm. H. Emory, U.S.A., under whose command part of the type material was col-
lected, "I take great pleasure, therefore, in dedicating this species to that distinguished officer."

(Agassiz)

66 bulletin: museum of comparative zoology

(Cibolo Creek, Boerne, Tex.); no. 7, July 15, 1926, p. 7 (Cedar Creek,
Henderson Co., Tex.); no. 19, 1929, p. 15 (Trinity Riv., Ft. Worth,
Tex.); no. 23, June, 1931, p. 16 (Colorado Riv., Trevis Co., Tex.);
Baylor Bull., 38, no. 3, Aug. 1935, p. 23 (Texas: Cibolo Creek), p. 32
(Texas: Real Co.).— ORTENBURGER, Proc. Oklahoma Acad. Sci.,
6, pt. 1, 1926, p. 100 (LeFlore Co., Oklahoma).— STRECKER and
WILLIAMS, Contr. Baylor Univ. Mus., no. 12, Dec. 27, 1927, pp.

11, 15 (San Marcos and Blanco Rivers, Texas).— RUST, Blatt.
Aquar. Terrarienk., 45, 1934, p.—, sep., p. 12.— LITTLE and KEL-
LER, Copeia, 1937, no. 4, Dec. 31, pp. 216, 221 (Mesilla Valley,
Dona Ana Co., New Mexico). — GAIGE, Univ. Michigan Stud. Sci.,

12, 1937, p. 304 (Mexico, Tamaulipas, Rio Purificaci6n, N. of Ciudad
Victoria).— DELLINGER and BLACK, Occas. Pap. Univ. Arkansas
Mus., no. 1, June 1938, p. 46 (? Arkansas: Salina Riv. near Benton?
[probably Texas, U.S.N.M. no. 17823-L.S.]).— SMITH, Ann. Car-
negie Mus. Pittsburgh, 27, 1939, p. 312 (Mexico: Tamaulipas;
Nuevo Laredo).

1870. — Aspidonectes emyda "Agassiz," GRAY, Suppl. Cat. Shield Rept. Brit.

Mus., pt. 1, p. 95 (lapsus).
1870. — Aspidonectes georgii "Agassiz," GRAY, Suppl. Cat. Shield Rept. Brit.

Mus., pt. 1, p. 109 (lapsus).

1893. Platypeltis emoryii BAUER, Proc. Amer. Philos. Soc, 31, p. 220 (emen-
dation).— Amyda emoryii STRECKER, Copeia, no. 162, 1927, p. 9
(food habits); Contr. Baylor Univ. Mus., no. 15, July 10, 1928, p. 6
(Bosque Riv. near Valley Mills, Bosque Co., Texas); no. 16, Aug. 4,
1928, p. 21 (Texas: vernacular names).

Agassiz (p. 407), as character aiding in identifying this species,
calls attention to skin of the carapace being "dotted all over with
small whitish tubercles like grains of sand." This is not a specific
character, it seems to be a condition of the skin due to season or age, as
similarly "sandpaper" specimens are encountered in several of the
species.

Skull. The skulls of A. emoryi and A. svinifera are very much
alike. The snout in emoryi is slightly shorter and somewhat broader
anteriorly, the nasal cavity relatively shorter and the angle formed by
the anterior processes of the prefrontal bones more obtuse. The
alveolar surface of the maxillaries are somewhat wider. In these re-
spects the emoryi are even closer to A. fero.v.

As the differences in the skull between A. spinifera and emoryi are
very slight, many skulls can hardly be told apart. In the former the
choanae and the intermaxillary foramen average a trifle larger. The
orbit in A. spinifera is also placed slightly more backward on the

STEJNEGER: AMERICAN SOFT-SHELL TURTLES 67

average than in A. emoryi. As a consequence the snout appears a
mere trifle longer. However, it is difficult to understand how Boulen-
ger (Cat. Chel. Brit. Mus., 18S9, pp. 245-246) came to diagnose A.
emoryi as having "the snout (on the skull) obtuse, hardly as long as the
diameter of the orbit," and the other two {spinifera and/< rox) having
it "a little longer." In 26 skulls measured by me the horizontal diame-
ter of the orbits in ferox, spinifera, and emoryi averages 20.0, 19.6, and
19.1 mm respectively and the snout (as measured from the orbit)
24.9, 26.5, and 24.4 mm. In the six emoryi measured by me the snout
is 2.5, 3.0, 4.0, 5.5, 1.5, and 2.0 mm longer than the ort.it. The relative
dimensions of snout and orbit are therefore unavailable as a diagnostic
character. It should be noted, finally, that in emoryi the alveolar
surface of the maxilla of medium sized skulls is slightly wider than in
the other two, but too slightly and variably so to be of much help in
diagnosing.

Plastron. Referring to Siebenrock's remarks about the plastron of
.4. emoryi (Sitz. Ber. Akad. Wiss. Wien, Math. Nat. KL, 111, 1902, p.
830) it is well to note that the bones are essentially as in A. spinifera.
The entoplastral angle in adults seems to be more acute. The callosi-
ties appear to be more developed in a male (U.S.N.M. no. 26426);
they are almost as large as in the A. mutica figured by him (p. 823, fig.
5), but of course without trace of callosities on entoplastron or epi-
plastra. The anterior portion of the epiplastra is rather long.

Color of live specimens

Two adult males from Houston, Texas, are colored as follows. Xo.
94335 has the top of head and dorsal aspect of neck dark olive green,
becoming gradually more green on posterior half of neck. Carapace,
bony disk Van Dyke brown, bistre on the soft parts, with irregular
blackish brown anastomozing spots. Sides of neck almost citron yellow
fading into whitish on the middle line of the ventral aspect of the neck
which anteriorly changes into verditer blue on the throat, darkening
to almost indigo blue on mentum and outer half of lower jaw and
tympanic region, and extending a little below and backwards on side
of neck; the yellow of the side of the neck fades into a dull olive yel-
low with a few scattered, almost blackish spots.

Tubercles on anterior edge of soft carapace large (3-4 mm.), tri-
angular, spaced apart by the width of their bases.

Underside white, also feet, but fingers and webs pale (dull) sage
green with obscure dark marblings.

68 bulletin: museum of comparative zoology

Their measurements differ as follows:

d* ad. 94335 & ad. 94336

Soft carapace, long 342 mm. 354 mm.

286 " 288 "

Height of body 86 " 84 "

Tip of tail beyond carapace 40 32

In No. 94336 the colors are essentially as in the other except that
on head and neck the yellow is a little deeper and that there are a
large number of small blackish spots on -sides of head, even including
lower eyelid, lips, and base of proboscis, and definite blackish lines
running from eye obliquely backward to base of lower jaw which
is also outlined by similar lines; dusky obscure large (average 15
mm.) ocellar markings on under side of neck. Tubercles like 94335.

Two specimens collected by Dr. and Mrs. A. H. Wright, U. S. N. M.
nos. 94456-7 are colored as follows. In no. 94456 the neck and iris
chrome yellow more or less pale, iris with horizontal black lens.
Carapace above uniform wood brown. Marginal edge washed with
chromium green. Upper soft parts of head and legs pale chromium
green washed with wood brown on top and sides of head. Tail above
white, with posterior central part dull chromium green washed with
cadmium. Tubercles on carapace pale buff, minute, round, densely
scattered over the whole carapace, hand, and soft parts. Underside
white. Neck underneath very pale cadmium shading anteriorly into
pale verditer blue washed with pale cadmium (on throat). Dark
markings on throat strongly tinged with bluish. Fleshy lips pale
cadmium yellow. Pale cadmium yellow on fork of snout fading away
anteriorly. Triangle, base between eyes slightly angular.

No. 94457 has the same data, but a dark ring of minute blackish
specks surrounding tubercles. Underside of foot same color as upper
chromium green parts — spotted.

stejneger: American soft-shell turtles

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70 bulletin: museum of comparative zoology

Another male, U.S.N.M. 104240, from Pecos River, near Dryden,
Terrell Co., Texas. F. M. Setzler, coll. shows still further variation.

Anterior border of disk very obscurely tuberculated. Skin on cara-
pace shagreened like bony disk. Skin on flaps smooth, leathery: a
round smooth area over each. The fontanelle smooth. Part of flap
behind bony disk with a regular pattern of whitish "pimples" or
tubercles, those along the middorsal line in pairs. No "sandpaper"
effect. Palms and soles yellowish white, unspotted. Carapace (day
or two after death) : flap Isabella color (Ridgway, pi. Ill, fig. 23); bony
disk more tawny olive (fig. 17); neck and limbs above pale olive gray
like pickled unripe olives, ventral half of neck yellowish white with a
faint trace of a similarly colored band from lower edge of eye back-
wards along the neck. All upper soft parts with numerous black dots
more or less arranged in longitudinal series.

Geographical distribution

Rivers of Texas, north into southern Oklahoma and Arkansas, west
(introduced?) to southeastern California, the adjacent portion of
Lower California, and Clark County, Nevada; east to western Louisi-
ana; northern Mexico.

List of specimens in the U. S. National Museum

7176

Okla., Old Ft. Cobb

E. Palmer May

4, 1868

7614-20,

Mex., Matamoras,

7662-25

Tamaulipas

L. B. Couch

7628-33

Mex., Matamoras,

Tamaulipas

7635-6

7637-8 juv.

Tex., Rio Bravo

A. Schott

7640 juv.

" Brazos R.

G. C. Shumard

7641 juv.

" El Paso del Norte

Dr. Webb

7642 cf adol.

" Brownsville

7644 juv.

n a

7700 (029536)

ad.

" New Braunfels

7701 juv.

" Rio Grande del

Norte

G. Wurdemann

7747 juv.

" Rio Seco

Capt. Pope

stejneger: American soft-shell turtles

7854 juv.

7855 juv.

Tex.

, Brownsville

Dr. Kennedy

(Cotype of
Aspidonedes
emoryi
Agassiz)

8925

J. C. Merrill

May, 1877

10789 juv.

San Antonio

C. W. Schuermann June 1879

17823 juv.

Ark.

, Benton

Jordan & Gilbert

19626-8

20846 9 ado!.

Tex.

, Ft. Hancock

E. A. M earns

Jan. 28, 1893

21408 juv.

26426 ad.

Tex.

, Ft. Clark

E. A. Mearns

26427 9 ad.

26428-36

45545

. tt

Boquillas

V. Bailey

May 25, 1901

46073

mouth of Devil's

River

W. Lloyd

Sept, 26, 1890

46074 juv.

Eagle Pass

Oct. 22, 1890

55601

McLennan Co.

J. Hurter

1906

66147 juv.

La.,

Madisonville

May 29, 1886

71627-8 adol.

Ariz

., Phoenix

V. Housholder

May 1, 1926

78515-6 juv.

Tex.

, Coleto Creek

J. D. Mitchell

Oct., 1905

78517

Tex.

, Guadalupe R.

J. D. Mitchell

Aug., 1912

83690

Christoval

C. E. Burt

Apr. 25, 1931

94335 d* ad.

near Houston

A. C. Chandler

94336 c? ad.

Houston

94456-7

Orange

A. H. Wright

Apr. 17, 1934

95386 c5 juv.

16 V2 mi. S.E. of

Caddo Lake

C. E. Burt

Apr. 1, 1934

95773 pull.

Llano River, Kimble

Co.

S. Mulaik

Aug. 10, 1933

100089 cf adol.

La.,

near Napoleonsvilk

: C. E. Burt

1935

100090 juv.

100380 d" adol.

Plaquemine

June 8, 1935

100419 <? adol.

Spanish Lake near

St,

Gabriel

June, 1935

103678 9

Tex.

, Boquillas

T. Smith

Aug. 6, 1937

104240 9 ad.

Pecos River near
Dryden

F. M. Setzler

72 bulletin: museum of comparative zoology

Amyda agassizii ' (Baur)
Plates 26-30

1857 .—Piatypeltis ferox AGASSIZ, Contr. Nat. Hist. United States, pt. 1, p.

401 (part) (not of Schneider).
Aspidonectes ferox COKER, North Carolina Geol. Surv., Bull. No. 14,

1906, p. 66 (South Carolina: Darlington Co., Society Hill, Peedee

River).— CORRINGTON, Copeia, No. 172, Nov. 15, 1929, p. 82

(Congaree Riv., between Columbia and New Brookland, Lexington

Co., South Carolina).
1888. Piatypeltis agassizii BAUR, Amer. Natural., 22, p. 1121 (type locality,

Georgia; type M.C.Z. no. 37172).
Pelodiscus agassizii BAUR, Proc. Amer. Philos. Soc, 31, July 1893,

p. 218.
Trionyx agassizii HAY, Indiana Geol. 17 Rep., 1892, p. 552; Batr.

Rept. Indiana, 1893, p. 144 (part, U.S.N.M. no. 8359).
1899. Aspidonectes agassizii JORDAN, Man. Vert. North Amer., ed. 8, p.

206 (part only: Ga.) (emendation).
1923. Trionyx spiniferus agassizii SIEBENROCK, Verh. Zool. Bot. Ver.

Wien, 73, Aug. 1923, p. 188 (part: Georgia).
1939.— Trionyx spiniferus POPE, Turtles U. S., Canada, pi. 45, figs. 98-99

(Broad River near Columbia, Richland Co., South Carolina) (not of

LeSueur).

The agassizii Group

In outward appearance Amyda agassizii differs very little from the
members of the spinifera group, but the skull distinguishes it at once
from all other American soft-shell turtles. In practically all the charac-
ters which differentiate it from them it agrees with the Asiatic mem-
bers of the group represented by the collective Amyda sinensis. A.
agassizii therefore may be treated as a member of a separate group,
thus more insistently emphasizing its isolated position and« insuring
the positive identification of specimens from its comparatively re-
stricted range in the United States.

Among Louis Agassiz's collection of colored drawings of turtles by
Burkhardt (mostly from life) are several painted in November and
December 1855, and inscribed as "Trionyx ferox, Ga. Dr. Daniel."
They are excellent pictures of Amyda agassizii, two of an "adult"
specimen (upper and lower surfaces) and one (upper side) of a young
specimen, but unfortunately they are without indication of size and
locality. They were sent to Agassiz by Dr. W. B. Daniel from Sa-
vannah, Ga. (Contr. Nat. Hist. U. S., 1, p. 401). The fact that they

1 Named for Louis Agassiz to indicate that the species was included by him in his account of
Piatypeltis ferox.

stejneger: American soft-shell turtles 73

came from the recorded type locality of T. ferox evidently influenced
Agassiz to regard these specimens as topo typical of ferox and hence
he recorded them as such.

The status of Baur's Platypeltis agassizii has never been fully ex-
plained. In 1888 (Amer. Natural., vol 22, p. 1121) the name appears
for the first time in the following sentence: " Platypeltis ferox of Agassiz
is not Testudo ferox Schneider, but a new species, which may be called
Platypeltis Agassizii." No further description or indication is given,
but evidently reference is intended to Agassiz's account of the species
in his Contribution to the Natural History of the United States, pt. 1 ,
pp. 401-403. A careful examination of Agassiz's text compared with
the material available to him at the time he wrote his account shows,
that his Platypeltis ferox is a mixture of two species due to the fact
that the ranges of both species overlap in Georgia and that the startling
color pattern of the very young specimens of the true Testudo ferox of
Schneider was unknown by him. Dr. Baur, in studying Agassiz's
material in the Museum of Comparative Zoology, designated the
specimen marked "Ferox Ga. No. 1" as the type of P. agassizii with
his own hand, and it must-be accepted as such.1 In 1893, after having
examined skulls and restudying the question of the generic relation-
ships of the Trionychids (Proc. Amer. Philos. Soc, vol. 31, July 1893,
p. 217) he came to the conclusion that his P. agassizii was not only
specifically but generically different from Schneider's P. ferox and re-
ferred it to Fitzinger's genus Pelodiscus with several Asiatic species.

In the meantime Dr. O. P. Hay, assuming that Agassiz's (op. cit.)
pi. 6, fig. 3, of his young Platypeltis ferox with the strongly marked two
black marginal rings represented the young P. agassizii (which has a
similar pattern), applied the name to the form which Agassiz had de-
scribed earlier as P. asper (Indiana Geol. 17 Rep., 1892, p. 552; Batr.
Rept. Indiana, 1893, Batr. Rept. Indiana, 1893, p. 144). This con-
fusion between A. agassizii and A. asper before their diagnostic fea-
tures were well understood was caused by the specimen U.S.N.M. No.
8359, a young A. agassizii, which was alluded to by Dr. Hay (op. cit.)—
though without giving the number — under A. agassizii as follows:
"This species belongs to the Southern States from South Carolina to
Texas. A single specimen has been forwarded to the National Museum
from Madison, Ind.". The specimen has the 2 black rings of the disk
characteristic of A. agassizii very plainly marked (pi. 30,) and as this
is also the normal pattern of the very young A. asper, Dr. Hay took

1 In a letter dated Feb. 13, 93 to Stejneger, he wrote: "The only Pelodiscus agassizii which I
have seen is at Cambridge Mus. (the type)."

bulletin: museum of comparative zoology

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stejneger: American soft-shell turtles

for granted that the agassizii and asper were synonymous, an error
followed by many subsequent authors.

The fact is that the locality "Madison, Ind." attributed to specimen
No. S359 in the original book of entry on June 25, 1875, is erroneous.
There is no record as to the donor or collector of the specimen and it
was entered on the book as part of an accumulation of miscellaneous
material. It is catalogued as Trionyx ferox and there is no original
label attached to the specimen, which only bears the tin-tag 8359, at
a time when the Museum had started a campaign to collect living
reptiles and amphibians to serve as models for the series of painted
casts in the exhibition series of the North American vertebrate fauna.
The tin-tagging and uncritical re-entry of many old specimens the
record of which was lost has been the source of many errors. (Note
reference to this specimen by Cahn in his Turtles of Illinois, 1937, p.
200, under Amyda ferox).

Geographical Distribution
Rivers of Georgia and South Carolina; North into southern North

Carolina.

List of specimens in U. S. National Museum

8359 juv.
8708 9 adol.
029034 9
30822 juv.
51981 (M.C.Z. 1598)

cf adol.
66859 9 adol.
71681 9 adol.

91282-3 ad. 9 ,

juv. d"
91310 9 adol.

91311-2 juv.

Ind., Madison ??
Ga., Milledgeville
Ga. ?

Ga., Baker Co.
)

Ga., Savannah
Ga., Augusta
S.C., Greenwood

T. H. Bean
R. Hessel
Brimley Bros.

W. B. Daniel
S. F. Hildebrand
Dr. Barrett,
A. L. Pickens

July, 1876
June 7, 1902

91491 juv.

91533

92521-3 J* adol.

92583 (M.C.Z.) adol. Ga.

92584 (M.C.Z. 1601)

9 Ga., Savannah

Ga., above Price Id.,

Savannah River E. H. Wood July, 193

S.C., 5 mi. W. of Plum

Branch, Savannah R. E. H. Wood
S.C., 5 mi. W. of Plum

Branch, Savannah R. " "

S.C., Batesburg L. Brodie

" Murray Lake C. E. Burt July 5, 1933

" Parks ville K. McNeill 1933

PLATES

PLATE 1

Stejnegeh — Soft-Shell Turtles

PLATE 1

Skull of Amy da (spinifera)

Fig. 1. View from above. Fig. 2. View from below. Fig. 3. View from left
side, including mandible. Fig. 4. View of mandible from above. Fig. 5. An-
terior plastral outlines of Amyda ferox U.S.N.M. 60496 (upper); Amyda
emoryi U.S.N.M. 94456 (middle); Amyda spinifera U.S.N.M. 101386 (lower).

alv. mb.

Alveolar surface of man-

pm.

premaxillary (intermaxil

dible

lary)

alv. mx.

Alveolar surface of max-

pof.

postfrontal (postorbital)

illa

prf.

prefrontal

art.

articular

pro.

prootic (otosphenoid)

boc.

basioccipital

pt.

pterygoid

bsp.

basisphenoid (para-

qj-

quadratojugal (para-

sphenoid)

quadratum)

cho.

choana

qu.

quadrate

cond.

occipital condyle

qu. art.

articulation of quadrat*

cor.

coronoid

with lower jaw

den.

dentary

s. ang.

supraangular

exoc.

exoccipital

soc.

supraoccipital

fr.

frontal

splen.

splenial

int. max.

for. intermaxillary foramen

sq.

squamosal

J"-

j«gal

sym.

symphysial

mx.

maxillary

temp, fossa

temporal fossa (inter

orb.

orbit

temporal foramen)

pa.

parietal

tymp. cav.

tympanic cavity

pal.

palatine

vom.

vomer

paoc.

paroccipital (opisthotic)

BULL. MUS COMP. ZOOL

-temp
fossa.

Stejneger. Soft-Shell Turtles. Plate 1.

pm.

~mt max for

alv. mx

temp,
fossa

Fig. 1

Fig. 2

pro
pot ju. Pil-MJ-

sopaoc

s.ar>

Fig. 4

Fig.

PLATE 2

Stejneger — Soft-Shell Turtles

PLATE 2
Amyda mutica 9, Fairport, Iowa. U.S.N.M. 53521.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 2.

■ « •

V <

, w
A ¥ A

<H-

3Sri

PLATE 3

Stejneger — Soft-Shell Turtles

PLATE 3

Upper view of Amyda mutica, U.S.N.M. 95186 from Medicine Lodge River,
near Lake City, Kansas. Also left side of head.

BULL MUS COMP ZOOL.

Stejneger. Soft-Shell Turtles. Plate 3.

PLATE 4

Stejneger — Soft-Shell Turtles

PLATE 4

Lower view of Amyda mutica, U.S.N.M. 95186 from Medicine Lodge River,
near Lake City, Kansas.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 4.

PLATE 5

Stejneger — Soft-Shell Turtles

PLATE 5

Skull of type of Amyda ferox in British Museum, from Georgia.

BULL MUS COMP. ZOOL.

Stejneqer. Soft-Shell Turtles. Plate 5.

PLATE 6

Stejneger — Soft-Shell Turtles

PLATE 6

Skulls of Amydaferox (U.S.N.M. 029475, 029459, 029464 and 029462 from
Kissimmee, Florida) showing extremes in the width of the alveolar surface and
in the outline of the maxilla.

BULL. MUS COMP ZOOL.

Stejneger. Sot-Shell Turtles. Plate 6.

PLATE 7

Stejneger — Soft-Shell Turtles

PLATE 7 .

Upper view of amyda ferox, U.S.N.M. 86828, Tamiami Trail near Birdon,
Florida.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 7.

PLATE 8

Stejneger — Soft-Shell Turtles

PLATE 8

Lower view of A my da- ferox, U.S.N.M. 86828, Tamiami Trail near Birdon,
Florida.

BULL. MUS. COMP. ZOOL.

Stejneger. Suet-Shell Turtles. Plate 8.

PLATE 9

Stejneger — Soft-Shell Turtles

PLATE 9

Amyda ferox, U.S.N. M. 86492 from Tamiami Trail about 15 miles from
Miami City, Florida.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 9.

PLATE 10

Stejneger — Soft-Shell Turtles

PLATE 10

Amydaferox, U.S.N.M. 60496 from Auburndale, Florida.

BULL. MUS COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 10.

PLATE 11

Stejneger — Soft-Shell Turtles

PLATE 11

Upper view of Amyda spinifera, U.S.N.M. 101386 from Middle Fork, Hol-
ston River, Seven Mile Ford, Virginia.

BULL. MUS. COMP. ZOOL

Stejneger. Soft-Shell Turtles. Plate 11.

" t«

Bg? '

PLATE 12

Stejneger — Soft-Shell Turtles

PLATE 12

Lower view of Amyda spinifera, U.S.N.M. 101386 from Middle Fork, Hol-
ston River, Seven Mile Ford, Virginia.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 12.

PLATE 13

Stejneger — Soft-Shell Turtles

PLATE 13

Head of Amyda spinifera, U.S.N.M. 101386 from Middle Fork, Holston
River, Seven Mile Ford, Virginia.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 13.

PLATE 14

Stejneger — Soft-Shell Turtles

PLATE 14

Cotype of Amyda nuchalis, Mus. Comp. Zool. 1623 from Cumberland River,
Tennessee. Upper view.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 14

zJs

PLATE 15

Stejneger — Soft-Shell Turtles

PLATE 15

Cotype of Amyda nuchalis, Mus. Comp. Zool. 1623 from Cumberland River,
Tennessee. Lower view.

BULL MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 15.

PLATE 16

Stejneqer — Soft-Shell Turtles

PLATE 16

Upper and lower views, and side of head of Amyda spinifera aspera. Cotype,
Mus. Comp. Zool. 1622 from Lake John, Florida.

BULL MUS COMP. ZOOL

Stejneger. So't-Shell Turtles. Plate 16.

PLATE 17

Stejneger — Soft-Shell Turtles

PLATE 17

Amyda spinifera aspera, U.S.N.M. 95191 from Canton, Mississippi. Upper
view.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 17.

PLATE 18

Stejnegeh — Soft-Shell Turtles

PLATE 18

Amyda spinifera aspera, U.S.N. M. 95191 from Canton, Mississippi. Lower
view.

BULL MUS COMP. ZOOL.

Stejneqer. Soft-Shell Turtles. Plate 18.

PLATE 19

Stejneger — Soft-Shell Turtles

PLATE 19

a. Amyda ferox juv., U.S.N.M. 61087 from Auburndale, Florida, b. Amy da
ferox juv., U.S.N.M. 84603 from Chesser's Island, Okefinokee Swamp, Georgia,
c. Amyda aspera, M.C.Z. 1608A from Mobile, Alabama.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 19.

r9H

PLATE 20

Stejneqer — Soft-Shell Turtles

PLATE. 20

Agassiz drawing of Amyda emoryi (upper view) which corresponds fairly
closely with the type (M.C.Z. 1910) collected at Brownsville, Texas by Col.
Emory.

BULL MUS. COMP. ZOOL

Stejneger. Soft-Shell Turtles. Plate 20.

^:$itik

.y i \ > -

PLATE 21

Stbjneger — Soft-Shell Turtles

PLATE 21

Agassiz drawing of Amyda emoryi (lower view) which corresponds fairly
closely with the type (M.C.Z. 1910) collected at Brownsville, Texas by Col.
Emory.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 21.

■&•>•■

PLATE 22

Stejneger — Soft-Shell Turtles

PLATE 22

Skull of Amyda emoryi, U.S.N.M. 78517, collected at Guadalupe River,
Victoria County, Texas by J. D. Mitchell.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 22.

Aft

PLATE 23

Stejneger — Soft-Shell Turtles

PLATE 23

Upper view of Amyda emoryi, U.S.N.M. 94336 from near Houston, Texas.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 23.

PLATE 24

Stejneger — Soft-Shell Turtles

PLATE 24
Lower view of Amyda emoryi, U.S.N. M. 94336 from near Houston, Texas.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 24.

PLATE 25

Stejneger — Soft-Shell Turtles

PLATE 25
Head of Amyda emoryi, U.S.N.M. 94336 from near Houston, Texas.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 25.

JL-.)k

PLATE 26

Stejneger — Soft-Shell Turtles

PLATE 26

Amyda agassizii Juv., U.S.N.M. 8359, wrongly recorded from Madison,
Indiana. Upper view.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 26.

i'l •

XSJii-

^flt&'i °

Q 0

•

PLATE 27

Stejneger — Soft-Shell Turtles

PLATE 27

Upper view of Amyda agassizii, U.S.N.M. 92521 from Parksville, South
Carolina.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 27.

■ >

1 i

hi,

PLATE 28

Stejneqer— Soft-Shell Turtles

PLATE 28

Lower view of Amyda agassizii, U.S.N.M. 92521 from Parksville, South
Carolina.

BULL MUS. COMP ZOOL

Stejneger. Soft-Shell Turtles. Plate 28.

PLATE 29

Stejneger — Soft-Shell Turtles

PLATE 29
Head of Amyda agassizii, U.S.N.M. 92521 from Parksville, South Carolina.

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 29.

PLATE 30

Stbjneger — Soft-Shell Turtles

PLATE 30

Skull of Amyda agassizii, Philadelphia Acad. Nat. Sci. 106 ( = 371).

BULL. MUS. COMP. ZOOL.

Stejneger. Soft-Shell Turtles. Plate 30.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 2

CONTRIBUTION TO THE ORNITHOLOGY OF THE
HAWAIIAN ISLANDS

By E. H. Bryan, Jr. and J. C. Greenway, Jr.

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

May, 1944

PUBLICATIONS
OF THE .

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
investigations by the Staff of the Museum or of reports by spec-
ialists upon the Museum collections or explorations.

Of the Bulletin, Vols. I to XCIII, and Vol. CXIV, No. 1
have appeared and of the Memoirs, Vol. I to LVI.

After 1941 no more Memoirs are to be published.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 2

CONTRIBUTION TO THE ORNITHOLOGY OF THE
HAWAIIAN ISLANDS

By E. H. Bryan, Jr. and J. C. Greenway, Jr.

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

May, 1944

No. 2 — Contribution to the Ornithology of the Hawaiian Islands '
By E. H. Bryan, Jr. and J. C. Greenway, Jr.

FOREWORD

For over a year prior to the entrance of the United States into the
war, Mr. E. H. Bryan, Jr., Curator of Collections at the B. P. Bishop
Museum, and Mr. J. C. Greenway, Associate Curator of Birds at the
Museum of Comparative Zoology, collaborated in the preparation of
a Check-List of Birds of the Hawaiian Islands. The finished draft of
the list was completed early in 1941, but, due to the fact that the
senior author was called to active duty as an army reserve officer,
his final comments were not received for inclusion until recently. In
the meantime the junior author was commissioned in the Navy; con-
sequently the final preparation of the Check-List for the printer
devolved upon the undersigned. No introduction had been prepared,
but I was so fortunate as to discover in the Bryan-Greenway corres-
pondence an outline of the history of Hawaiian ornithology which,
though not prepared with such a purpose in mind, nevertheless appears
suitable as such. A brief abstract has previously appeared in the
Proceedings Hawaiian Academy of Science for 1935 (Special Publica-
tion no. 26, Bernice P. Bishop Museum) p. 5-6.

Shortly before being ordered to active duty Greenway had incor-
porated his studies of the Hawaiian drepanids in a rearrangement of
the genera of this most interesting family; with the publication of this
Check-List, the opportunity to publish this account seems opportune.

Ornithologists all wish for the safe and speedy return to their chosen
fields of Capt. Edwin H. Bryan, Jr. A. U.S. and Lieut. James C.
Greenway, Jr. U.S.N.R.

James L. Peters

Cambridge, Mass.
23 June 1943

1 Published with the aid of a special gift from Mr. G. R. Agassiz.

80 bulletin: museum of comparative zoology

HAWAIIAN BIRDS1

By E. H. Bryan, Jr.

Curator of Collections, B. P. Bishop Museum2

In 1778, when the third voyage of Captain Cook made contact
between Hawaii and the so-called civilized world, there lived in Hawaii
about 100 different kinds of birds. Most of these were well known to
the Hawaiians, each kind being called by a native name. A few species
were eaten; the feathers of several kinds were used to decorate the
kahili or royal standards of the kings and high chiefs ; feathers of the
mamo, oo, iiwi, and to a less extent those of the apapane and o-u,
were tied in decorative patterns upon fine mesh network of olona
fiber to form royal uniforms and ceremonial robes; and in one way or
another many found a place in native life and lore.

Although Captain Cook did not live to see his native land again,
and although there was no trained naturalist with this third voyage,
his companions seem to have procured specimens of about sixteen
species of Hawaiian birds, and to have carried them safely back to
ornithologists in Europe. These first native Hawaiian species to be
made known to science included the iiwi, mamo, apapane, amakihi,
akepa, akialoa, o-u, oo, thrush, elepaio, rail, crow and migrant golden
plover. A specimen of the oo-aa, was also taken, but this was not
recognized as distinct from the oo, until 1855.

The iiwi has the distinction of being the first native Hawaiian
species of bird to be technically described. Barthold Lohmann, who
sailed with Cook's last expedition, brought specimens of it to Cassel,
Germany, where it was described by Georg Forster, in 1780 as Certkia
coccinea. During the next four years John Latham gave brief descrip-
tions and popular names to nine other species in his General Synopsis
of Birds. In 1788, Gmelin gave scientific names to eleven, including
these, largely on the basis of Latham's notes. Six species can be recog-
nized from the names and descriptions given by James King in his
account of Cook's voyage.

Thus did Hawaiian ornithology get away to a good start, only to
sink into quiescent repose until toward the end of the 19th century.
This came about partly through neglect, but to quite a large extent
because of a series of unfortunate circumstances.

1 Address of the retiring president, Hawaiian Academy of Science, May 18, 1935.

2 Published by permission of the Director, B. P. Bishop Museum.

BRYAN AND GREE.WVAY: HAWAIIAN' BIRDS 81

In the first place, the types of these early species, which had been
placed in the British Museum, by some mischance became lost or
destroyed. This led to considerable confusion in later technical sum-
maries, catalogs and lists, because of the lack of authentic specimens
to which to refer. In the second place, the naturalists with Vancouver,
Kotzebue, and other early expeditions which touched Hawaii, seem
to have either quite neglected the interesting bird life, or else to have
made collections and observations which resulted in no publications
for the advancement of science.

In 1825, H. M. S. 'Blonde', under command of George Anson,
seventh Lord Byron, carried back to Hawaii the bodies of Kame-
hameha II and his queen who had died in England. On board was
Chaplain Richard R. Bloxam and his brother Andrew, an enthusiastic
young naturalist. Ornithologists of the day hoped that young Andrew
Bloxam might get some of the curious Hawaiian birds and produce
an interesting publication about them. He apparently got very little
official encouragement, in spite of which he obtained specimens of nine
species of land birds on Oahu, including the now extinct thrush. He
worked hard over his specimens, and placed them, all properly labelled,
at the disposal of the Lords of the Admiralty. The scientific "Appendix
to the Voyage of the Blonde", published in 1826, is said to have been
edited bv a woman who had onlv a few of Bloxam's notes to guide her.
This, combined with some poor judgment on the part of the ornitho-
logical gentlemen of the British Museum, who identified the specimens,
made the results, as Professor Alfred Newton put it1 "a disgrace to
all concerned, since, so far from advancing the knowledge of the
subject, it introduced so much confusion as to mislead many sub-
sequent writers," especially in the absence of the specimens, which
disappeared not long after.

A few years later another good opportunity to advance Hawaiian
ornithology was missed. Dr. J. K. Townsend, the American traveller,
and the well-known naturalist, Thomas Nuttall, made a trip together
to Hawaii. Arriving in January, 1836, they spent three months col-
lecting on Oahu and Kauai. Townsend, returning at the end of the
year, found the Prussian naturalist, Herr Deppe, at Honolulu, and
with him spent a few months in the pursuit of natural history, leaving
Hawaii in March, 1837. A few of Deppe's birds were described by
Lichtenstein, and most of Townsend's specimens have been carefully
preserved, the bulk of them in the Academy of Natural Sciences,

1 Alfred Newton, Ornithology of the Sandwich Islands, Nature, 45, p. 466, 1892.

82 bulletin: museum of comparative zoology

Philadelphia. Had Townsend only published a list of his species,
and had both men but made a scientific record of their observations,
the knowledge of Hawaiian bird life would have been greatly advanced.

In the course of a six months' stay on Hawaii in 1840, the enthus-
iastic collectors, Peale and Pickering, of the United States Exploring
Expedition, obtained a large collection of birds. Most of these speci-
mens were lost in the shipwreck of the "Peacock," one of the ships of
Commodore Wilkes' squadron. Still another misfortune occurred in
1848. Peale's report on the birds of the group was just off the press,
and only a few copies had been distributed, when the entire stock was
destroyed by a fire. It is the opinion of some ornithologists that John
Cassin's new edition of this report, published ten years later, was no
improvement on Peale's original work.

In 1852 Dr. Hartlaub wrote a review of Peale's lost work, and later
he compiled the first list of Hawaiian birds, published in 1854. He
listed but 30 species of birds (of which five have not been accepted).
Of the remaining 25, only 16 are land birds, and only 14 are perching
birds.

It remained for our own Sanford B. Dole to produce an extensive
list of the Hawaiian birds. This was published first in 1869 in the
Proceedings of the Boston Society of Natural History, and later,
with additions and corrections, in the Hawaiian Annual for 1879.
Judge Dole listed 53 species, of which four were described as new.

Although in the vicinity of Hawaii from July 27 to August 19, 1875,
the scientific party of H. M. S. 'Challenger' collected only 24 bird
skins, of which only one species was new to science, the Hawaiian
duck, described in 1878 by P. L. Sclater, as Anas wyviUiana.

Revival of Interest

Considerable credit for a revival of interest in Hawaiian birds
should go to Professor Alfred Newton, of Cambridge University,
from whose interesting account of early Hawaiian ornithology some
of the foregoing has been condensed. Because of Dr. Newton's enthu-
siasm, two young Cambridge naturalists undertook collecting which
led to magnificent contributions to Hawaiian ornithology.

The first of these was Scott B. Wilson who went to Hawaii and
collected birds at Professor Newton's request. Leaving Liverpool on
February 24, 1887, he arrived in Honolulu on April 8, having made
a brief visit at Washington, D. C, with Dr. Leonhard Stejneger,
who had done considerable work on Hawaiian birds, especially those

BRYAN* AND GKKKNWAY: HAWAIIAN BIRDS 83

sent to him at the U. S. National Museum by Valdamar Knudsen of
Kauai. Wilson stayed on the islands until the close of 1888, taking
back with him to England a large and valuable collection of bird
specimens, rich in new species. This collection formed the basis for
the beautifully illustrated "Aves Hawaiienses," issued in sections
from 1890 to 1899. Wilson again visited Hawaii, in 1896, without
adding much to scientific knowledge. Besides collecting skins, he
carefully preserved whole specimens of the birds in alcohol. Upon
these specimens Dr. Hans Gadow based anatomical studies which
led to some new and startling conclusions as to the relationships of
many Hawaiian species.

Meanwhile, the Hon. Walter Rothschild, the other naturalist
interested by Professor Newton at Cambridge, sent his collector,
Henry Palmer, to the islands to procure specimens and data. Arriving
in December, 1890, Palmer collected on nearly all the main islands
of the group in company with George C. Munro, and also made a trip
to Laysan and Midway Islands, stopping at, or at least sighting,
most of the small rocky islets and reefs en route. By August, 1893,
when he left Honolulu, Palmer had collected 1832 bird specimens,
including all but seven of the species previously known, and in addi-
tion fifteen which Rothschild described as newT to science.

Rothschild produced the elaborate "Avifauna of Laysan and the
neighbouring islands with a complete history to date of the birds of
the Hawaiian possessions," published in London in three parts, which
appeared August, 1893, November 1893, and December 1900. These
contain over 300 large, well printed, folio pages, and numerous photo-
graphs and artistic colored plates drawn by Keulemans and Frohawk.
The work enumerates 116 species of birds, giving valuable notes on
food, habits, eggs, nests, and distribution, as w7ell as descriptions.

In 1898 Dr. H. Schauinsland spent three months on Laysan, and
in his fascinating little book, printed in Bremen in 1899, he lists the
birds which he found there. In 1900 he also furnished notes on a few
birds of Molokai.

In 1892 R. C. L. Perkins came to Hawaii under the joint auspices
of the Royal Society of London and the British Association for the
Advancement of Science, writh financial assistance from Bernice P.
Bishop Museum. For a decade he made extensive collections of
Hawaiian birds, as well as of insects and other land animals. His
section of the 'Fauna Hawaiiensis' on the birds was published Novem-
ber 1903. It gives an interesting and valuable summary of their dis-
tribution and relationships, especially of the native perching birds.

84 bulletin: museum of comparative zoology

In 1901, there appeared a "Key to the birds of the Hawaiian group"
by William Alanson Bryan, followed by a number of other lists and
notes on the birds of different islands of the group, written by him
and by Alvin Seale, published by Bernice P. Bishop Museum. Also,
in 1902-3, there was printed "A complete list of the birds of the
Hawaiian possessions, with notes on their habits," by H. W. Henshaw.
All of these publications have helped to complete and bring up to date
our knowledge of Hawaiian ornithology.

By 1930 the large number of birds which had been imported from
many parts of the world prompted the production of a report by
E. L. Caum on the "Exotic birds of Hawaii," published by B. P.
Bishop Museum in 1933.

Recent articles on Hawaiian birds include: — a report on sea bird
conditions on Laysan in 1911 by Homer Dill and W. A. Bryan; four
chapters in W. A. Bryan's Natural History of Hawaii; a popular
account of bird life in the Hawaiian Islands Bird Reservation by
Alexander Wetmore; four chapters in Hawaiian Nature Notes by E. H.
Bryan, Jr. ; and various articles in journals and the daily press.

Summary of Hawaiian Birds

With this brief historical background, let us consider the different
groups of birds which are found in Hawaii, and some of the interesting
facts concerning them. The list of the birds reported as having been
found in Hawaii, outside of captivity, both native and introduced,
now numbers 232 species (see table 1.) Of these, 77 species might be
considered as endemic; 18 are sea birds which nest regularly in the
archipelago, but which have a wider range; 1 species, the night heron,
is indigenous, but is found elsewhere; 8 are regular winter migrants;
34 have been recorded from time to time, and might be classed as
occasional migrants or chance arrivals; and 94 are immigrant exotic
species. Of these introduced species, 53 are probably established, and
41 are probably not established, although the exact status of several
is difficult to determine. So little field work has been done during the
past 30 years that it is not wise to state what species are really extinct.

Without reference to taxonomic classification, most of the Hawaiian
birds may be divided into three fairly well defined groups: (1) the
birds of the mountain forests; (2) the sea birds and migrants; and
(3) the introduced species. A few lowland species, such as the two
birds of prey, the black-crowned night heron, the native duck and

BRYAN AND GRKENWAY: HAWAIIAN BIRDS

goose, the extinct rail, the mud hens, the stilt, and the peculiar land
birds of Laysan, now all but extinct on that island, which do not fall
into these three general groups, might be given separate consideration.

Table 1. NUMERICAL SUMMARY OF HAWAIIAN BIRDS

Family

Endemic

Indigenous

& wide-
ranging sea

Regular

Migrants

< •(■(visional

& chance

arrivals

Tnt
Estab.

roduced

Not Estab

Diomedeidae

Procellariidae

Phaethontidae

Sulidae

Phalacrocoracidae

Fregatidae
Ardeidae

1
1

Threskiornithidae

Phoenicopteridae
Anatidae

Aceipitridae
Falconidae

Cracidae

Tetraonidae

Phasianidae

16?

Numididae

Meleagrididae
Turnicidae

1
1

Gruidae

1 ?

Rallidae

Charadriidae

Recurvirostridae

Phalaropodidae
Laridae

2
9

Columbidae

Psittacidae

Strigidae
Alcedinidae

Alaudidae

Corvidae

Paridae

Timaliidae

Mimidae

1 ?

Turdidae

Sylviidae

Museicapidae

86 bulletin: museum of comparative zoology

Prionopidae

Sturnidae

Zosteropidae

Drepanididae

Meliphagidae

Ploceidae

Icteridae

Fringillidae

19 8

Total Native

104

Total Introduced

Grand Total

232.

The Sea Birds

1 ?

34 53 41

Concerning the sea birds, found throughout the Hawaiian group,
little need be said here. Most of them are species widespread in the
Pacific, and although scientists argue over their scientific names they
are fairly easy to recognize. They include: the two large albatross,
the thieving frigate, three species of homely boobies, five kinds of
noisy terns, the burrowing shearwaters and petrels, and the graceful
white-and-red-tailed tropic birds. Most of them stay well away from
man's habitations. Not that they are afraid of him. Rather, man
and sea birds are rivals for a large and steady supply of fresh fish,
without which an extensive bird colony could not exist.

President Theodore Roosevelt and other thoughtful citizens have
helped to protect our sea birds by establishing for them the Hawaiian
Islands Bird Reservation on islands and reefs to the northwest of
Kauai. Here they are not only kept from molestation, but also from
time to time efforts are made to keep their rather poor environment
from becoming any worse, through occasional visits to kill off rabbits
and plant more vegetation.

Migratory Birds

Each year there come to Hawaii large numbers of tourists, concern-
ing whom no record is kept by our efficient Tourist Bureau. They
spend no money here, but they do a very considerable amount of good.
They are the migratory birds. Arriving from a more than 2,000 mile
flight, thin and hungry, they alight on our beaches, fields and pastures
and make short work of great numbers of caterpillars, grasshoppers

BRYAN AND GBEENWA1 : HAWAIIAN BIRDS 87

and other insects, much to the benefit of agriculture. They arrive and
depart with great regularity, coming in the late summer or early fall
and leaving again in the spring, to return to nest in their cold northern
home. Included among these gentle and helpful visitors, which should
be given a better welcome than the hunters' gun, are the golden
plover, the turnstone„ wandering tatler, sanderling, curlew, various
wild ducks, and several others.

The native birds of Hawaii's lowlands, marshes, ponds, valleys,
and grassy slopes are becoming very rare. The flightless rail is gone;
the duck and goose are making a last valiant stand against extermina-
tion; the mud hens, stilts, and night heron seem to be holding their
own. One still sees Pueo, the native owl, winging his low flight above
the tops of guava bushes and across grassy hillsides at sunset, in search
of rats and lizards. But Io, the native hawk, is scarce in Hawaii, due
to the mistaken idea that it was an enemy to the poultrymen.

Birds of the Mountain Forests

The birds of the mountain forests are all native perching birds and
include 50 to 55 endemic species of that order. Twenty-three of the
twenty-four genera in this group are also endemic, only Corpus being
found elsewhere in the world. These undoubtedly make up the most
interesting part of the Hawaiian bird fauna. Whence did they come?
What were their ancestral relationships? Why are there so many
genera and species, even one whole family, found nowhere else in the
world? Why are they so rapidly becoming extinct? These are ques-
tions which interest not only the ornithologist, but also the zoologist,
the biologist, the conservation expert, and the student of evolution.
I can only try to hint at the answers to some of these questions.

These native perching birds, found in the forests of the main islands
of the group, include: the crow, found only in the Kona and Kau
districts of Hawaii; six different species of thrushes, five of which
(found on five different islands) are closely related, and one, on Kauai,
somewhat different in form, food, and habits; three closely related
species of elepaio or flycatcher, on three different islands; four species
of oo, on four different islands, and one other very distinct honey-eater
of Hawaii; and between 40 and 45 different species of native "honey
creepers," belonging to a family found nowhere else in the world.
I give this range of number for the drepanids because various authori-
ties differ in their concepts of specific differences.

88 bulletin: museum of comparative zoology

Native Hawaiian Honey Creepers

The drepanids themselves are worthy of extensive study. There
are, let us say, forty species, representing eighteen genera. These
have very diverse appearance; some are black and golden, some red
and black, some green and yellow; some have shprt bills for picking up
insects, some long, slender bills for sipping nectar from the base of
the long-tubed blossoms of the native lobelias, and some have heavy,
almost finch-like or parrot-like beaks for cracking seeds. Only three
species, the iiwi, apapane, and o-u, are found on more than two islands.
Yet it is believed, that all are the descendents of a common ancestor,
or at most two ancestral stocks.

The careful anatomical studies of Dr. Hans Gadow suggest that
their nearest relatives are the Neotropical and Central American
Coerebidae. Peter P. Suskin (International Ornith. Kongress 1926,
Verhandl., VI, pp. 375-381, 1929) suggests descent from Fringillidae
of the Malaysian region. We can, perhaps, imagine that many thou-
sands of years ago there reached Hawaii from far away tropical America
or southeastern Asia a little group of these birds. A pair of them,
at least, survived, and establishing themselves, became the common
ancestors of our interesting native family. We can but suppose that
their descendents, becoming isolated on different islands of the group,
with different kinds of environment and different sources of food
supply, developed into the diverse species which we find today, or
rather, in many cases, — yesterday, for a number are very rare, even
extinct.

Origin of Hawaiian Bird Groups

The same explanation as to origin might be offered for other groups
of native perching birds. For example, one ancestral kind of immigrant
flycatcher, arriving not so many centuries ago, has given rise to the
three species of elepaio, one each on Hawaii, Oahu, and Kauai, which
look so very much alike that they might better be considered only
geographic races, rather than different species. One immigrant an-
cestral form may have been the progenitor of the four kinds of oo,
the Oahu species of which has not been seen alive since 1837; another
was the ancestor of Chaetoptila, a Hawaii honey-eater, now probably
extinct. The thrushes descended from two ancestral stocks; one
ancestor giving rise to one species each of A-Maui on Kauai, Oahu,
Molokai, Lanai, and Hawaii, and the little, weevil-eating Puaichi

BRYAN AND GREENWAY: HAWAIIAN BIRDS 89

having descended from another. Thus, all of the native Hawaiian
perching birds might have developed, as Dr. Perkins has pointed out1
"from at least six and not more than seven successfully immigrant
species."

It would make a fascinating study, by tracing out the distribution
of the nearest relatives of these various groups of birds, to try to
determine whence they came; and, by studying their degree of develop-
ment and evolution, as well as their local distribution, to attempt to
guess how long they have been in Hawaii. One interesting point
is the fact that, although the distances between the Hawaiian Islands
are not very great, and although the native forest birds have fairly
good powers of flight, their distribution is so definitely limited and
restricted.

Why Native Species are Becoming Extinct

One hears various explanations as to why the native species of
Hawaiian birds are so rapidly becoming rare or extinct. We are told
by those who have not made a study of the subject that they were
killed off by the native feather-gatherers or chased away by the minah
birds, or their eggs destroyed by the mongoose. These, in certain
instances, may have been contributing causes, but they have certainly
not been the basic reason for the changes which have come over the
native forest life. The main reason may be stated in these few words:
Nature established a balance in Hawaii, and Nature's balance has
been upset. What is this "Balanee in Nature" and what has caused
its upset?

Long ago, before the coming of man with his ships, new plants and
animals arrived from foreign parts and established themselves in
Hawaii only at long, infrequent intervals of time. This was because
of Hawaii's extreme isolation — more than two thousand miles from
the regions from which the ancestors of most of the native species
came. The kinds which managed to arrive and gain a foothold had to
adjust themselves to their new home. If they were in harmony with
their new environment they survived; if they were not in harmony,
they either became adjusted or modified until they were, or else they
perished. There have been many types of environment in HawTaii:
bare lava flows, hot dry deserts, fertile valleys, marshy lowlands,
grassy hills, wet forests, summit bogs, bare, snow-capped peaks.

1 R. C. L. Perkins, Fauna Hawaiiensis, vol. I, pt. iv, p. 369, 1903.

90 bulletin: museum of comparative zoology

Each offered a habitat for a different type of life, or a reason for
modification or adjustment. In each different environment a balance
was established, with the ironclad policy, "fit in or perish." If a
species became over-abundant it was brought back to a proper numer-
ical balance either through lack of food supply, or by the activity of
its enemies, which took advantage of its abundance to multiply them-
selves, only in turn to be starved or eaten back to normal.

Upset of Nature's Balance

Then into this nicely balanced island world came man, with his
foreign plants and animals, his thoughtless or ruthless waste, his
clearing and burning. The nice balance of Nature was upset. Cattle,
sheep, pigs, and goats ran wild through the forests, so damaging the
undergrowth that it no longer protected the roots of the growing trees.
Rain eroded away the soil; seeds could no longer germinate; and both
forest trees and undergrowth died off. Their places were quickly
taken by foreign plants which no longer furnished food or protection
to native insects and birds. Whole associated groups, interdependent
upon each other, were disassociated. Foreign birds were imported,
bringing with them diseases against which the specialized native
birds had no immunity. With food and shelter destroyed, and danger
threatening them in the lowlands, the native birds either sought new
homes in the depths of the forests, where they in turn upset nicely
adjusted Nature, or else they became extinct.

Foreign Birds

And now, because native birds are scarce, the cry is for more and
yet more foreign birds. Some kinds have been introduced to satisfy
the desire of the hunter; pheasants from the mountains of Mongolia
or Java; grouse and quail from northwestern America, China, Japan
and Australia; partridges from the slopes of the Himalayas; doves
and pigeons from Panama, India, China, Australia, and the Malay
Isles. Some, like the skylark and cardinal, have been brought because
of sentiment or to satisfy esthetic taste. Some have been introduced
for a more useful purpose, such as the horn fly-eating willie wag-tail,
the meadow lark, even the much maligned, but very useful minah.
A few accidental escapes we could well get along without.

BRYAN' AND GREENWAY: HAWAIIAN BIRDS 91

Fortunately for Hawaii, our Territory's scientific- advisers have
advocated a policy of rigid protection for our native and useful bird
life, and a strong stand against the promiscuous importation of foreign
birds and animals. This need not include birds which are to be kept
in a proper aviary, carefully safeguarded against their escape. I can
see much that is good in recent plans for extensive breeding and
exhibition of showy and interesting birds in Honolulu, so long as they
are not allowed to go wild. But I cannot approve of wholesale intro-
duction and liberation, without adequate study by experts as to the
consequences.

You may say, now that Nature's balance has been so badly upset,
and Hawaii's avian heritage is passing out of the picture, what harm
can a little more upset do ! But the birds may not be the only part of
our natural history to be effected by such introductions. Forestry,
agriculture, and all branches of animal life are so interrelated that
one cannot tell, without considerable study, what far reaching con-
sequences may result from one foolish introduction. We have effective
quarantine laws to protect Hawaii against insect pests and plant
diseases, which we all admit help tremendously to safeguard our
island paradise. Let us try to protect what bird life we have left, and
also try to avoid the risk of further upset of Nature's balance, just
for the sake of adding a flash of color or a chirp of song.

92 bulletin: museum of comparative zoology

' CHECK-LIST OF THE BIRDS
OF THE HAWAIIAN ISLANDS

By E. H. Bryan, Jr. and J. C. Greenway, Jr.

Part I. NATIVE BIRDS

Order PROCELLARIIFORMES

Family DIOMEDEIDAE— Albatrosses

Genus Diomedea — Albatrosses

Diomedea Linnaeus, Syst. Naturae, ed. 10, 1, 1758, p. 132. Type, by subse-
quent designation, D. exulans Linn.

Diomedea nigripes Audubon

Diomedea nigripes Audubon, Ornith. Biog., 5, 1839, p. 327. (Pacific Ocean,

lat. 30° 44' N., long. 146° W.)
Black-footed Albatross, Brown Gooney
Phoebastria nigripes reischekia Mathews, Bull. British Ornith. Club, 51, 1930,

p. 29. (New Zealand.)
Diomedea chinensis Temm. in Rothschild, Avifauna of Laysan, etc., p. 55

(corrected p. 292).
Diomedea albatrus (chinensis) in Wilson and Evans, Aves Hawaiiensis, p. xxv.

Range. North Pacific, breeding on islands northwest of Hawaii;
Marshall Islands and Bonin Islands.

Recorded in Hawaiian Group from Midway (breeding), Pearl and
Hermes Reef (sight), Lisianski, Laysan (breeding), French Frigate
Shoal (breeding) (sight), Necker (sight), "Bird Rock" (Nihoa) (sight),
Kaula (breeding), Lehua (sight) (Caum). Many sight records at sea.
"This is the Gooney that follows in the wrake of ships from San Fran-
cisco to Honolulu and Hilo" (Henshaw).

Nests behind the beaches on the periphery of the islands. A hybrid,
D. nigripes x D. immutabilis, from Midway is in the Bishop Museum.

Diomedea immutabilis Rothschild

Diomeda immutabilis Rothschild, Bull. British Ornith. Club, 1, 1893, p. xlviii.

(Laysan Island.)
Gooney, White Albatross, Laysan Island Albatross

Range. Central and North Pacific.

Breeds on Laysan and Midway Islands. Nests in interior of the
islands. To be seen at sea from islands off the coast Lower California
to the Bonin Islands.

BRYAN AND GREENWAY : HAWAIIAN BIRDS 93

Recorded in the Hawaiian Group from Midway (breeding), Laysan
(breeding), Lisianski (breeding), Necker Island (breeding), Pearl and
Hermes Reef (breeding), Gardner Island, "Bird Island" (Nihoa)
(breeding). "Apparently common throughout Hawaiian main group,
though it is not known to breed on any of them" (Henshaw).

Family PROCELLARIIDAE

Genus Puffinus — Shearwaters

Puffinus Brisson, Ornithologie, 1, 1760, p. 56; 6, p. 130. Type, by tautonomy,
Puffinus Brisson = ProceUaria puffinus Briinnich.

Puffinus pacificus cuneatus Salvin

Puffinus cuneatus Salvin, Ibis, 1888, p. 353. (Krusenstern Island.)

Wedge Tailed Shearwater, Uau Kane, Uwau

Puffinus pacificus laysani Mathews, Birds of Australia, vol. 2, 1912, p. 83.

(Laysan Island.)
Puffinus knudseni Stejneger, Proceed. U. S. Nat. Museum, 11, 1888, p. 93.

(Kauai.)

Range. From about 140° west latitude (two days from San Fran-
cisco) to the Bonin Islands and south to the Equator.

Recorded in the Hawaiian Group from Midway, Pearl and Hermes
Reef (sight, Galstoff), Lisianski, Laysan (breeding), French Frigate
Shoal, Xecker (breeding), "Bird Island" (Nihoa), Kaula, Lehua,
Kauai, Oahu (breeding on Mokumanu, Manana and Popoia Islets)
(Munro).

Puffinus nativitatis Streets

Puffinus (Nectris) nativitatis Streets, Bull. U. S. Nat. Museum, 1877, no. 7

p. 29. (Christmas Island, Pacific Ocean.)
Black Shearwater, Christmas Island Shearwater

Range. Pacific Ocean from the vicinity of 25° north latitude south
to the vicinity of 25° south latitude.

Recorded in the Hawaiian Group from Midway, Laysan (breeding),
French Frigate Shoal (breeding), "Bird Island" (Nihoa) (breeding?),
Lisianski (Munro ms.), Gardner (sight record by Dr. Isenbeck re-
corded by Kittlitz 1834), Oahu (Mokulua Islet) (sight by Donagho),
Mokumanu Islet (breeding) (Munro).

Puffinus newelli Henshaw
Puffinus newelli Henshaw, Auk, 17, 1900, p. 246. (Waihee Valley, "Ulani" in

error, Maui Island, see Henshaw, Birds of Hawaiian Possessions, p. 117,

footnote.)
Newell' s Shearwater, Ao

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Range. Hawaiian Islands.

Recorded in the Hawaiian Group from ?Niihau, Kauai (breeding),
Molokai, Maui. We can find no records for Lanai.

Genus Pterodroma — Petrels

Pterodroma Bonaparte, Comptes Rendus Academie Sciences, Paris, 42, 1856,
p. 768. Type, by subsequent designation, Procellaria macroptera A. Smith.

Pterodroma phaeopygia sandwichensis (Ridgway)

Oestrelala sandwichensis Ridgway, Baird, Brewer and Ridgway's Water

Birds of North America, vol. 2, 1884, p. 395. (Hawaiian Islands.)
Dark-rumped Petrel, Uau or Uwau
Oestrelala phaeopyga In Wilson and Evans, Aves Hawaiiensis.

Range. Hawaiian Islands.

Recorded in the Hawaiian Group from Hawaii (breeding on Mauna
Kea and Mauna Loa), Kauai (breeding), Molokai (breeding) (eggs in
Bishop Museum), Lanai (Munro).

Henshaw (1902) found that the mongoose had driven these birds
from their nesting grounds on Hawaii.

Pterodroma leucoptera hypoleuca (Salvin)

Oestrelata hypoleuca Salvin, Ibis, 1888, p. 359. (Krusenstern Island, North

Pacific Ocean.)
Salvin' s White Breasted Petrel, Bonin Island Petrel

Range. Northern Pacific Ocean. Breeds on the Bonin Islands ?
and Hawaiian Islands.

Recorded in the Hawaiian Group from Midway (breeding — eggs
collected by T. M. Blackman), Laysan (breeding) (Rothschild),
(W. K. Fisher 1903), ? Kaula (Caum), Lanai (Munro).

Mathews (1934) records Pterodroma leucoptera longirostris Stejneger
(1893) as the breeding bird of the Bonin Islands. This is not the case.
Birds of the Bonin Islands differ from Japanese birds and agree with
Hawaiian birds in having grayer under wing coverts and in their
large size.

Krusenstern Rock probably does not exist, reported by Captain
Lisianski in lat. 22° 15' N., long. 175° 37' W., it has been dropped
from the Pacific Islands Pilot of the British Admiralty (ed. 1931)
and though it is recorded in the Hawaiian Islands Pilot of the U. S.
Hydrographic Office (ed. 1933), there has been no actual report of a
rock in this vicinity since 1804 when breakers were reported thirty
miles south of the supposed position of the rock. The type locality
of this bird is therefore doubtful.

BRYAN AND GREENWAY : HAWAIIAN BIRDS 95

Mathews (1934) records the breeding range of this bird as follows:
"Hawaiian Group (Laysan, Pearl, Hermes [sic], Lisianski, Krusenstern
Reef, Midway and Ocean' Islands and French Frigate Shoal)." We
cannot find that this bird has ever been recorded as breeding on any
island except Laysan.

Genus Bulweria — Petrels

Bulweria Bonaparte, Nuova Annales Sci. Nat. Bologna, 8, 1842, p. 426. Type,
by monotypy, Procellaria bulweri Jardine and Selby.

Bulweria bulwerii (Jardine and Selby)

Procellaria bulwerii Jardine and Selby, Illust. Ornith. 2, 1828, pi. 65. (Ma-
deira.)

Soft Nosed Petrel, Bulwer's Petrel

Bulweria bulweri pacifica Mathews and Iredale, Ibis, 1915, p. 607. (Imojima,
Bonin Islands.)

Bulweria anjinho Heineken in Wilson and Evans, Aves Hawaiiensis, p. 211.

Range. "Breeding on islands off the coast of China; the Bonin
Islands, Vulcan Islands, the western Hawaiians and the Marquesas
Islands in the Pacific Ocean; Madeira, the Salvages, Canary and
Cape Verde Islands in the Atlantic" (Peters).

Recorded in the Hawaiian Group from Laysan, French Frigate
Shoal (breeding), Necker Island (breeding), Kauai, Hilo, Hawaii
(Henshaw), Oahu (skin in Bishop Museum, G. C. Munro coll.).

W. A. Bryan does not distinguish between sight records and col-
lected specimens. His records are from Necker and Bird Island
(Xihoa).

We quite agree with Hartert (Novitat. Zool., 33, 1926, p. 326) that
pacifica cannot be maintained.

Family HYDROBATIDAE

Genus Oceanodroma — Petrels

Oceanodroma Reichenbach, Aves Systema Naturae, 1852 (1853 fide Peters)
p. iv. Type, by original designation, Procellaria furcata Gmelin.

Oceanodroma castro cryptoleucura (Ridgway)
Cymochorea cryptoleucura Ridgway, Proceed. U. S. Nat. Museum, 4, 1882,

p. 337. (Kauai, Hawaiian Islands.)
Hawaiian Stormy Petrel, Ake-Ake (Henshaw), Oeoe (W. A. Bryan)

Range. Hawaiian Islands.

Recorded in the Hawaiian Group from Midway (Bartsch), French
Frigate Shoal, Niihau, Kauai.

"The natives report a small petrel, which they call by the above

96 bulletin: museum of comparative zoology

name (Ake-Ake), as common on the fishing grounds five or ten miles
off the coast of Hawaii, and I have little doubt that it is this species
..." (Henshaw).

OCEANODROMA MARKHAMI TRISTRAMI Salvin

Oceanodroma tristrami Salvin, Catalogue Birds British Museum, 25, 1896,

p. 354. (Sendai Bay, Japan.)
Sooty Petrel, Tristram's Petrel, Fork Tailed Petrel.
Oceanodroma fuliginosa (Gmelin) in Rothschild, Avifauna of Laysan, etc.,

p. 308; W. A. Bryan, Birds of the Hawaiian Group, p. 13 (footnote);

Henshaw, Birds of the Hawaiian Islands, p. 119; Oceanodroma tristrami

Stejneger in Bartsch, Auk, 34, 1922, p. 486; W. K. Fisher, Bull. U. S. Fish

Commission, 23, 1923, p. 795.

Range. From Japan south and west to the Hawaiian Islands.

Recorded in the Hawaiian Group from Midway (Bartsch), Laysan
(breeding), ?"Bird Island" (Nihoa) (W. K. Fisher), Lanai (skin in
Bishop Museum, G. C. Munro coll.).

In our opinion, Salvin's description of tristrami is identifiable and
Hartert's arguments (Vogel palaarktischen Fauna, p. 1416) have
no force under the Rules of Zoological Nomenclature. Mathews
records Cymochorea (markhami) owstoni Mathews and Iredale 1915,
as the breeding bird of the Hawaiian Islands in the belief that the
name tristrami is not identifiable.

Order PELECANIFORMES

Family PHAETHONTIDAE

Genus Phaethon — Tropic or Bos'n Birds

Phaethon Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 134. Type, by sub,
sequent designation, Phaethon adherens Linn. (Gray, List Genera Birds-
1840, p. 80). For synonymy see Peters, Check-List of Birds of the World,
1, p. 77.

Phaethon rubricauda rothschildi (Mathews)

Scaeophaethon rubricauda rothschildi Mathews, Birds Australia, 4, 1915, p. 303.

(Laysan and Niihau.)
Red-tailed Tropic Bird, Bos'n Bird, Koaeula
Scaeophaethon rubricauda brevirostris Mathews, Birds Australia, 4, 1915,

p. 303. (Bonin Islands.)
Phaethon rubricauda Boddaert in Rothschild, Avifauna of Laysan, etc., pp. 33,

294, 296 and other works.

BRYAN AND GKEKXWAY: HAWAIIAN BIRDS 97

Range. Pacific Ocean from the Bonin and Marianas Islands to the
Hawaiian Islands.

Recorded in the Hawaiian. Group from Midway (breeding; eggs in
Bishop Museum, T. M. Blackman coll.), Laysan (breeding), French
Frigate Shoal (sight by W. K. Fisher), Necker, Kaula, Lehua, Niihau,
Mokumanu Islet, Oahu (sight, Munro), Lanai (sight, Munro).

Wilson and Evans record this bird breeding on Kauai and Niihau,
and occurring on Hawaii.

Phaethon lepturus dorotheae Mathews

Phaethon lepturus dorotheae Mathews, Austral Avian Rec, 2, 1913, p. 7-

(Queensland.)
White Tailed Tropic Bird, Salmon Tailed Tropic Bird, Bos'n Bird, Koae,

Haakoae
Phaethon lepturus Daudin in Rothschild, Avifauna of Laysan, etc., p. 296;

Henshaw, Birds Hawaiian Islands, p. 114, and other works.
Phaethon aethereus Bloxam (nee Linn.) in Wilson and Evans, Aves Hawaiiensis,

p. 186.

Range. Australia and certain Pacific islands.

Recorded in the Hawaiian Group from Kauai, Oahu, Maui, Molokai
(specimens in Mus. Comp. Zool.), Hawaii.

Hartert (Novitat. Zoologicae, 32, 1925, p. 276) remarks that
dorotheae is probably not distinct from lepturus, which is the breeding
bird of Mauritius. This does not appear to us to be the case. The
species is in need of revision.

Family SULIDAE

Genus Sula — Boobies

Sula Brisson, Ornithologie, 1, 1760, p. 60; 6, p. 494. Type, by tautonomy,
"Sula" = Sula leucogaster Boddaert.

Sula sula rubripes Gould

Sula rubripes Gould, Synopsis Birds Australia, pt. 4, 1838, append., p. 7.

(New South Wales = Raine Island, N. Queensland, fide Mathews.)
Tied Footed Booby

Sula piscatrix Linn, in Rothschild, Avifauna of Laysan, etc., pp. 27, 297.
Sula piscator Linn, in Wilson and Evans, Aves Hawaiiensis, p. xxv; Henshaw,

Birds Hawaiian Islands, p. 113; W. A. Bryan, Key, Birds Hawaiian

Group, p. 15.

98 bulletin: museum of comparative zoology

Range. "Islands of the Indian and tropical western and central
Pacific Oceans" (Peters).

Recorded in the Hawaiian Group from Midway, Lisianski and Laysan
(breeding), French Frigate Shoal (Bryan), "Bird Island" (Nihoa)
(breeding), Lehua (Rothschild), Niihau, but see W. K. Fisher, Bull.
U. S. Fish Comm., 23, 1903, p. 797, Kaula, Kauai, Oahu (on Mokumanu
Island) (breeding), and many sight records at sea.

Sula leucogaster plotus (Forster)

Pelecanus plotus Forster, Descriptiones Animalium, etc., (ed. Lichtenstein)

1844, p. 278. (Near New Caledonia.)
Booby, Common Brown Booby, Brown Vested Booby
Sula sula Linn, in Rothschild, Avifauna of Laysan, etc., pp. 28, 297; Wilson

and Evans, Aves Hawaiiensis, p. xxv; Henshaw, Birds Hawaiian Islands,

p. 114; W. A. Bryan, Key, Birds Hawaiian Islands, p. 15; Henshaw,

Birds Hawaiian Possessions, p. 114.

Range. "Western and central tropical Pacific to northeastern
Australia" (Peters).

Recorded in the Hawaiian Group from Midway, Lisianski and Laysan
Islands, French Frigate Shoal (sight) (Fisher), Necker (breeding),
"Bird Island "(Nihoa) (breeding), Kaula (Caum), and many sight
records at sea off Niihau, Oahu (Mokumanu Islet) (Munro).

Sula dactylatra person at a Gould

Sida personata Gould, Proc. Zool. Soc. London, 1846, p. 21. (North and
northeast coasts of Australia = Raine Island, North Queensland, fide
Mathews.)

Booby, Blue Faced Booby, Masked Booby, Masked Gannet

Sula cyanops Sundevall in Rothschild, Avifauna of Laysan, etc., p. 25; Wilson
and Evans, Aves Hawaiiensis, p. xxv; Henshaw, Birds Hawaiian Islands,
p. 113; W. A. Bryan, Key, Birds Hawaiian Group, p. 15.

Range. "Breeds on islands of the central and western tropical
Pacific Ocean" (Peters).

Recorded in the Hawaiian Group from Midway (breeding), Lisianski,
Laysan (breeding), French Frigate Shoal (breeding), Necker (breed-
ing), "Bird Island" (Nihoa) (breeding), Kaula, Lehua (Caum).

Family PHALACROCORACIDAE

Genus Phalacrocorax — Cormorants

Phalacrocorax Brisson, Ornithologie, 1, 1760, p. 60. Type, by tautonomy,
Phalacrocorax = Pelecanus carbo Linnaeus.

BRYAN AND GREENWAY: HAWAIIAN BIRDS 99

Phalacrocorax pelagicus pelagic is Pallas

Phalacrocorax pelagicus Pallas, Zoographia Rosso-Asiatica, 2, 1811, p. 303.

(Eastern Kamchatka and the Aleutian Islands.)
Pelegic Cormorant

Range. Breeds on the coasts of northeastern Asia, Aleutian Islands
to south-central Alaska; south in winter to Japan and Puget Sound.

Recorded in the Hawaiian Group {as an accidental visitor) from
Laysan 1896 (Rothschild), ?Hilo, Hawaii 1900 (recorded by Henshaw
with doubt).

Family FREGATIDAE

Genus Fregata — Frigate Birds or Man o'War Birds

Fregata minor palmerstoni (Gmelin)

Pelecanns palmerstoni Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 573.

(Palmerston Island = Palmerston Atoll, Cook Group.)
Man o'War Bird, Ima
Fregata aquila Linn, in Rothschild, Avifauna of Laysan, etc., pp. 22, 297 and

other works.

Range. "Hawaiian Islands south to New Zealand" (Peters).

Recorded in the Hawaiian Group from Midway (breeding), Pearl and
Hermes Reef, Laysan (breeding), French Frigate Shoal, Necker
(breeding), "Bird Island" (Nihoa) (breeding), Kaula, Lehua, Kauai,
Oahu.

Order CICONIIFORMES

Family ARDEIDAE— Herons

Genus Nycticorax — Night Herons

Nycticorax T. Forster, Synoptic Catalog British Birds, 1817, p. 59. Type, by
tautonomy and monotypy, Nycticorax infaustus Forster = Ardea nycti-
corax Linn.

Nycticorax nycticorax hoactli (Gmelin)

Ardea hoactli Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 630. (In Novae

Hispania lacubus = Valley of Mexico.)
Black Crowned Night Heron, Auku Aukuu, Auku Kahili or Auku Kahili
Nycticorax nycticorax naevius (Boddaert) in Rothschild, Avifauna of Laysan,

etc., p. 265.
Nycticorax griseits (Boddaert) in Wilson and Evans, Aves Hawaiiensis, p. 201,

and Henshaw, Birds Hawaiian Islands, p. 102.

100 bulletin: museum of comparative zoology

Range. Northern United States and northeastern Canada south
through Mexico and Central America and east coast of South America
to eastern Argentine.

Recorded in the Hawaiian Group from Midway (skin in Bishop
Museum — D. R. Chisholm coll.), Oahu, Molokai, Hawaii — "all
islands" (Rothschild, AYilson and Evans, Henshaw).

The records of Demigretta sacra seem to be doubtful. As Bryan
(1901, p. 21) remarks, Dole's record of 1879 might just as well refer
to the Auku.

The identity of the White Heron which Dr. Finsch saw so long ago
at Kahalui is problematical.

Family THRESKIORNITHIDAE

Genus Plegadis — Glossy Ibises

Plegadis Kaup, Skizzirte Entwickelungs-Geschichte Natiirliches System, etc.,
1829, p. 82. Type, by monotypy, Tantalus falcinellus Linnaeus.

Plegadis guarauna (Linnaeus)

Scolovax gaurauna Linnaeus, Systema Naturae, ed. 12, 1, 1766, p. 242.

(Brazil.)
White Faced Glossy Ibis

Range. Northwestern North America south to Argentina.
Recorded in the Hawaiian Group (as an accidental visitor) from
Kauai, ?Maui (Henshaw), Molokai (Munro).

Order ANSERIFORMES

Family ANATIDAE

Genus Chen — Snow Geese

Chen Boie, Isis von Oken, 10, 1822, column 563. Type, by monotypy, Anser
hyperboreus Pallas.

Chen hyperborea hyperborea (Pallas)

Anser hyperboreus Pallas, Spicilegia Zoologica, etc., 1767 (1769?), fascicule 6

1769, p. 31. (Northeastern Siberia.)
Lesser Snow Goose

BRYAN AM) GKKK.WVAY: HAWAIIAN BIRDS 101

Range. Arctic North America south in winter to temperate zone.
Probably also Siberia south in winter to Japan.

Recorded in the Hawaiian Group (as an accidental visitor) from
Maui by Rothschild who received a specimen collected there by
Brother Matthias Newell; Oahu (J. E. Whitney coll.).

Genus Axser — Geese

A user Brisson, Ornithologie, 1, 1760, p. 58. Type, by tautonomy, Anser
domesticus = Anser anser Linnaeus.

Anser albifrons gambelli Hartlaub

Anser gambelli Hartlaub, Revue et Magasin Zoologie, etc., 1852, p. 7. (Texas

and southern United States.)
American White-Fronted Goose

Range. Northwestern North America south in winter to northern
California.

Recorded in the Hawaiian Group (as an accidental visitor) from
"a lake on Mr. Clark's estate at Honokaohau," Hawaii, where it was
shot December 18, 1891 (Rothschild), Molokai (G. C. Munro coll.).

Genus Philacte — Emperor Goose

Philacte Bannister, Proceedings Acad. Nat. Sci. Philadelphia, 1870, p. 131.
Type, by monotypy, Anas canagica Sewastianov.

Philacte canagica (Sewastianov)

Anas canagica Sewastianov, Nova Acta Acad. Sci. Imp. St. Petersburg, 13,

1802, p. 349, pi. 10. (Kanaga Island, Aleutian Islands.)
Emperor Goose

Range. Breeds on coasts of Siberia and Alaska, winters chiefly
in the Aleutian Islands.

Recorded in the Hawaiian Group (as an accidental visitor) from
Puna, Hawaii (Henshaw 1903).

Genus Branta — Brant or Brent Geese

Branta Scopoli, Annus I, Historico-Naturalis, 1769, p. 67. Type, by sub-
sequent designation, Anas bernicla Linnaeus (Bannister, Proceedings
Acad. Nat. Sci., Philadelphia, 1870, p. 131).

102 bulletin: museum of comparative zoology

Branta bernicla nigricans (Lawrence)

Anser nigricans Lawrence, Annals Lyceum Nat. Hist. New York, 4, 1846,

p. 171, pi. 12. (Egg Harbor, New Jersey.)
Black Brant

Range. Arctic coasts of Siberia and Arctic North America; south
in winter to Japan and North China, and to Lower California.

Recorded in the Hawaiian Group (as an accidental visitor) from Maui
(Rothschild).

Branta canadensis minima Ridgway

Branta minima Ridgway, Proceedings U. S. Nat. Museum, 8, 1885, p. 22.

("Pacific coasts of North America"; type from St. Michael's, Alaska.)
Cackling Goose
Bernicla Munroii Rothschild, Annals and Magazine Nat. Hist. (6), 10, 1892,

p. 108. (Kauai.)

Range. Breeds on the Bering Sea coast of Alaska and the Aleutian
Islands; in winter, west of the Rocky Mountains from southern
British Columbia to southern California.

Recorded in the Hawaiian Group (as an occasional or chance visitor)
from Kauai (Rothschild), ?Hawaii (Henshaw, q. v.), Molokai (G. C.
Munro coll.).

Genus Nesochen — Hawaiian Goose

Nesochen Salvadori, Catalogue Birds British Mus., 27, 1895, pp. 81, 126.
Type, by original designation and monotypy, Anser sandvicensis Vigors.

Nesocehn sandvicensis (Vigors)

Anser sandvicensis Vigors, List Animals Garden Zool. Soc, ed. 3, 1833, p. 4.
(Hawaiian Islands.)

Hawaiian Goose, Nene

Bernicla sandvicensis in Wilson and Evans, Aves Hawaiiensis, p. 187; Hen-
shaw, Birds Hawaiian Islands, p. 103.

Range. Hawaiian Islands.

Recorded in the Hawaiian Group from Hawaii. (Reports from
Maui, Kauai and Niihau have never been confirmed.) Breeds on the
Kona coast, and above 5000 feet; from the Kona District to the north-
east side of Mauna Kea, descending to 1000 feet and sea level in
winter. Nests in lava flows or open fields. Now much reduced in
numbers, though still breeding in a wild state as well as in captivity.

BRYAN AND GREENWAY: HAWAIIAN BIRDS 103

Genus Anas

Anas Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 122. Type, by subsequent
designation, Anas boschas Linnaeus = Anas platyrhynchos Linnaeus.

Anas platyrhynchos platyrhynchos Linnaeus

Anas platyrhynchos Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 125.

(Europe, type locality restricted to Sweden.)
Mallard

Range. Temperate zones of Europe, Asia and North America.
Recorded in the Hawaiian. Group (as an accidental visitor) from
Laysan (Schauinsland), Oahu, Molokai (Perkins).

Anas wyvilliana wyvilliana Sclater

Anas wyvilliana Sclater, Proc. Zool. Soc. London, 1878, p. 350. (Hawaiian

Islands.)
Hawaiian Duck, Koloa Maoli
Anas aberti Ridgway, Proc. U. S. Nat. Mus., 1, 1878, p. 250. (Mazatlan,

Mexico.)

Range. Hawaiian Islands. Recorded from Niihau, Kauai, Oahu,
Molokai, Maui and Hawaii in ponds and lakes along the coast and
in mountain streams and bogs to 8000 feet.

Recorded recently from Kauai, Mokulua Islets, N. E. coast of
Oahu (breeding) (Munro), Molokai, highlands of Hawaii (Baldwin).

The type of Anas aberti Ridgway was probably an escaped captive.
Mazatlan was formerly a port for vessels from China and possessed a
large bird market.

Anas wyvilliana laysanensis Rothschild

Anas laysanensis Rothschild, Bull. British Ornith. Club, 1, 1892, p. xvii.

(Laysan Island.)
Laysan Teal

Range. Laysan. Recorded by Kittlitz (Museum Senckenbergianum
1834, p. 124) from Lisianski but never confirmed.

Twenty birds left in 1923 (Wetmore, Nat'l Geographic Mag., 48,
no. 1, 1925, p. 103). Nine left in 1936 (W. F. Coultas, in Hit).

104 bulletin: museum of comparative zoology

Anas crecca carolinensis Gmelin

Anas carolinensis Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 533. (Carolina to

Hudson's Bay.)
Green Winged Teal

Range. Northern North America; in winter south to West Indies
and Panama.

Recorded in the Hawaiian Group (as an accidental visitor) from
Laysan (Schauinsland), Maui (G. P. Wilder coll.), Oahu (Northwood),
Molokai (G. C. Munro coll.).

Anas acuta acuta Linnaeus

Anas acuta Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 126. (Europe —

type locality restricted to Sweden.)
Pintail, Koloa Mapu

Range. Europe, Northwestern North America east to Iowa; south
in winter to West Indies, Panama and the Hawaiian Islands.

Recorded in the Hawaiian Group (as a regular migrant in winter)
from Laysan, Kauai, Oahu (Northwood), Hawaii. "Most of the
islands; common" (Perkins). "Well known to sportsmen as a winter
visitor" (Henshaw).

Genus Mareca — W7idgeons

Mareca Stephens, in Shaw, General Zoology, 12, pt. 2, 1824, p. 30. Type, by
subsequent designation, Mareca fistularis Stephens = Anas penelope
Linnaeus (Eyton, Monograph Anatidae, 1838, p. 33).

Mareca Americana (Gmelin)

Anas americana Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 526. (Louisiana

and New York.)
American Widgeon, Baldpate

Range. Northwestern to north-central North America; south in
winter to the Atlantic coast, West Indies and Central America.

Recorded in the Hawaiian Group (as an accidental visitor in winter)
from Laysan (Schauinsland), Oahu (Northwood), Maui (Perkins),
Molokai (Munro coll.).

BRYAN AND GRKKN WAY : HAWAIIAN' KIRDS 105

Genus Chaulelasmus — Gadwall Ducks

Chaulelasmus Bonaparte, Geographical and Comparative List of Birds of
Europe and North America, 1838, p. 56. Type, by monotypy, Anas
strepera Linnaeus.

Chaulelasmus streperus (Linnaeus)

Anas strepera Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 125. (Europe —

type locality restricted to Sweden.)
Gad wall

Range. Worldwide within the northern hemisphere.
Recorded in the Hawaiian Group (as an accidental visitor in winter)
from Oahu (Perkins), Molokai (Munro coll.).

Genus Spatula — Shoveller Ducks

Spatula Boie, Isis von Oken, 1822, col. 564. Type, by monotypy, Anas
clypeata Linnaeus.

Spatula clypeata (Linnaeus)

Anas clypeata Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 124. (Coasts of

Europe; type locality restricted to southern Sweden.)
Shoveller, North West Duck, Koloa Moha

Range. Worldwide within the northern hemisphere.
Recorded in the Hawaiian Group (as a regular migrant in winter)
from Laysan, Kauai, Oahu, Molokai, Lanai, Maui, Hawaii.

Genus Bucephala — Golden Eyes, Buffle Head

Bucephala Baird, Reports, Explorations and Surveys for a Railroad from the
Mississippi River to the Pacific Ocean, 9, pt. 2, pp. L, 788, 785. Type, by
original designation, Anas albeola Linnaeus.

Bucephala albeola (Linnaeus)

Anas albeola Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 124. (America =

Newfoundland ex Edwards.)
Bvffle Head

Range. Northwestern North America; in winter eastward and
southward in the United States.

Recorded in the Hawaiian Group (as an accidental visitor) from
Laysan (Rothschild), Oahu (Northwood), Maui (Perkins).

106 bulletin: museum of comparative zoology

Genus Nyroca — Diving Ducks

Nyroca Fleming, Philosophy of Zool. etc., 2, 1822, p. 260. Type, by tautonomy,
Anas nyroca Giildenstadt.

Nyroca affinis (Eyton)

Fuligula affinis Eyton, Monograph Anatidae, 1838, p. 157. (North America.)
Lesser Scaup Duck

Range. Northern North America; south in winter to Central
America.

Recorded in the Hawaiian Group (as an accidental visitor) from
Lanai (G. C. Munro coll.).

Genus Histrionicus — Harlequin Ducks

Histrionicus Lesson, Manuel d'Ornithologie, 2, 1828, p. 415. Type, by original
designation, Anas histrionica Linnaeus.

Histrionicus histrionicus pacificus W. S. Brooks

Histrionicus histrionicus pacificus W. S. Brooks, Bull. Mus. Comp. Zool.,

Cambridge, Mass., 59, 1915, p. 393. (Cape Shipunski, Kamchatka.)
Western Harlequin Duck

Range. Eastern Siberia and northwestern North America ; south in
winter to Japan and California.

Recorded in the Hawaiian Group (as an accidental visitor) from Lay-
san (P. E. H. Bompke coll. 1906).

Genus Mergus — Mergansers

Mergus Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 129. Type, by subse-
quent designation, Mergus castor Linnaeus = Mergus serrator Linnaeus.

Mergus serrator Linnaeus

Mergus serrator Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 129. (Europe.
Type locality restricted to Sweden.)

Range. Worldwide in the northern hemisphere ; south in winter to
temperate zone.

Recorded in the Hawaiian Group (as an accidental visitor) from 0§hu
(Bryan), Hawaii (Henshaw).

BRYAX AND GRKK.WVAY: HAWAIIAN BIRDS 107

Order FALCOXIFORMES
Family ACCIPITRIDAE

Genus Buteo — Soaring Hawks and Buzzards

Buteo Lacepede, Discours . . . Tableaux methodiques mammiferes . . •
Oiseaux, 1799, p. 4. Type, by tautonom}', Falco buteo Linnaeus.

Buteo solitarius Peale

Buteo solitarius Peale, U. S. Exploring Expedition, 8, 1848, p. 62. (Island of

Hawaii.)
Hawaiian Hawk, Io
Accipiter hawaii Dole, Hawaiian Almanac for 1879 (1878) p. 43.

Range. Island of Hawaii (all parts).

Genus Circus — Marsh Hawks

Circus Lacepede. Discours . . . Tableaux methodiques mammiferes oiseaux,
1799, p. 4. Type, by subsequent designation, Falco aeruginosus Linnaeus
(Lesson, Manuel d'Ornith., 1, 1828, p. 105).

Circus cyaneus hudsonius (Linnaeus)

Falco hudsonius Linnaeus, Systema Naturae, ed. 12, 1, 1766, p. 128. (Hudson
Bay ex Edwards.)

Marsh Hawk-
Range. Northern and central North America; south in winter to

Central America.

Recorded in the Hawaiian Group (as an accidental visitor) from Oahu

(Wilson).

Genus Pandion — Ospreys

Pandion Savigny, Description d'Egypt, Oiseaux, 1809, p. 69. Type, by
monotypy, Pandion fluvialis Savigny = Falco haliaetus Linnaeus.

Pandion haliaetus carolinensis (Gmelin)

Falco carolinensis Gmelin, Systema Naturae, 1, pt. 1, 1788, p. 263. (No type

locality. Carolina ex Catesby et al.)
American Osprey

108 bulletin: museum of comparative zoology

Range. Northern North America; south in winter to the West
Indies and South America.

Recorded in the Hawaiian Group (as an accidental winter visitor)
from ?Niihau (Dole), Oahu (Perkins), ?Molokai (Dole), ?Hawaii
(Dole).

Family RALLIDAE— Rails

Genus Porzanula

Porzanula Frohawk, Annals and Magazine Nat. Hist., London, (6), 9, 1892,
p. 247. Type, by monotypy, Porzanula palmeri Frohawk.

Porzanula palmeri Frohawk

Porzanula Palmeri Frohawk, Annals and Magazine Nat. Hist., London (6), 9,

1892, p. 247. (Laysan Island.)
Laysan Rail, Laysan Crake

Range. Laysan Island, where but two remained in 1923 (Wetmore
1925); Midway Island, where introduced in the latter part of the 19th
century (Rothschild, p. xiii) and where common (Denig et al).

Genus Pennula

Pennula Dole, Hawaiian Almanac and Annual for 1879 (1878), p. 54. Type,
by monotypy, Pennula millei (sic) Dole.

Pennula millsi Dole

Pennula millei [sic] Dole, Hawaiian Almanac and Annual for 1879 (1878),
p. 54. (Hawaii.)

Hawaiian Rail, Moho

Rallus ecaudotus (sic) King, in Cook's Voyage to the Pacific Ocean, 3, 1784,
p. 119. (Hawaiian Islands); Pennula ecaudata (King) in Wilson and
Evans, Aves Hawaiiensis, p. 171, and in Perkins, Fauna Hawaiiensis, 1,
pt. 4, p. 453, and older works. This name is preoccupied by Rallus ecau-
datus Miller 1783.

Range. Formerly Island of Hawaii in open grassy country or low
scrub just below the heavy rain forest, possibly in the Olaa district,
certainly on the windward side of Kilauea and perhaps about forty
miles north along the coast. ?Molokai (Perkins). Extinct, last speci-
men seen 1884 (Rothschild), ?1893 (Henshaw). Five specimens in
museums.

BRYAN AND GREENWAY: HAWAIIAN BIRDS 109

Pennula sandwichensis (Gmelin)

Rallus sandwichensis Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 717. ("in
insulis Sandwich," based on Latham's Sandwich Rail.)

Spotted Hawaiian Rail

Rallus obscurus Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 718. ("in insulis
Sandwich," based on Latham's Dusky Rail, and other authors.)

Pennula wilsoni Finsch, Notes Leyden Mus., 20, 1898, p. 77. Hawaiian
Islands.)

Range. Unknown. Now extinct. Known only from the unique
type in Leyden, Holland. Paler than millsi with central black spot on
feathers of back.

Genus Gallinula — Coots, Gallinules

Gallinula Brisson, Ornithologie, 1, 1760, p. 2. Type, by tautonomy, Gallinula
Brisson = Fulica chloropus Linnaeus.

Gallinula chloropus sandvicensis Streets

Gallinula sandvicensis Streets, Ibis, 1877, p. 25. (Oahu, Hawaiian Islands.)
Hawaiian Gallinule, Alae, Alae Ula

Range. Hawaiian Islands. Recorded from Kauai, Oahu, Molokai,
Maui, Hawaii in ponds, rice fields and taro patches.

Genus Fulica — Coots

Fulica Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 152. Type, by tau-
tonomy, Fulica atra Linnaeus.

Fulica Americana alai Peale

Fulica alai Peale, United States Exploring Expedition, 8, 1848, p. 224.

(Hawaiian Islands.)
Hawaiian Coot, Alae Keokeo

Range. Hawaiian Islands. Recorded from Niihau, Kauai, Oahu>
Molokai, Maui, Hawaii.

Order CHARADRIIFORMES

Family CHARADRIIDAE

Genus Squatarola — Plovers

Squatarola Cuvier, Regne Animal, 1, 1817 (1816), p. 467. Type, by tautonomy,
Tringa squatarola Linnaeus.

110 bulletin: museum of comparative zoology

Squatarola squatarola (Linnaeus)

Tringa squatarola Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 149. (Europe.

Type locality restricted to Sweden by Hartert, Vogel palaarktischen

Fauna, 2, p. 1553.)
Black-bellied Plover, Beetle Head

Range. Circumpolar in the arctic; south in winter to South Africa*
India, Australia, western South America.

Recorded in the Hawaiian Group (as an accidental visitor on migration)
from Oahu (Northwood et al), Hawaii (Henshaw).

Genus Pluvialis

Pluvialis Brisson, Ornitholigie, 1760, 1, p. 46; 5, p. 42. Type, by tautonomy,
Pluvialis aurea Brisson = Charadrius pluvialis Linnaeus = Charadrius
apricarius Linnaeus.

Pluvialis dominica fulva (Gmelin)

Charadrius fulvus Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 687. (Tahiti.)

Pacific Golden Plover, Kolea

Charadrius fulvus in Rothschild and other works.

Range. Siberia and Alaska; south in winter to India, Australia,
Indo-China, Pacific Islands.

Recorded in the Hawaiian Group (as a regular winter visitor) from
Midway, Laysan, Lisianski, French Frigate Shoal, Bird Island (Nihoa),
Niihau, Kauai, Oahu, Molokai, Maui, Hawaii, "all islands" (Perkins).
Arrive in August or September; depart in April or May as a rule, but
sometimes they remain during the summer, though probably they
never breed.

Genus Charadrius — Plovers

Charadrius Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 150. Type, by
tautonomy, Charadrius hiaticula Linnaeus.

Charadrius vociferus vociferus Linnaeus

Charadrius vociferus Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 150.

(North America = South Carolina ex Catesby.)
Killdeer

Range. Arctic and temperate zones and portions of the eastern
tropical; south in winter to northern South America.

Recorded in the Hawaiian Group (as an accidental visitor) from Maui
(W. A .Bryan).

BRYAN AND GREENWAY: HAWAIIAN BIRDS 111

Family SCOLOPACIDAE

Genus Numenius — Curlews

Numenius Brisson, Ornithologie, 17G0, 1, p. 48; 5, p. 311. Type, by tautonomy,
X miliums Brisson = Scolopax arquata Linnaeus.

Numenius tahitiensis (Gmelin)

Scolopax tahitiensis Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 656. (Tahiti

ex Latham.)
Bristle Thighed Curlew, Kioea

Range. Alaska; south in winter to the Pacific islands.

Recorded in the Hawaiian Group (as a regular winter visitor) from
.Midway, Laysan, Niihau, Kauai, Oahu, Molokai, Lanai (sight)
(Rothschild), Maui, Hawaii. Like the Golden Plover (Kolea), this
bird leaves in April or May and arrives in August or September, but
sometimes remains during the summer, though it probably never
breeds on the islands.

Genus Limosa — Godwits

Limosa Brisson, Ornithologie, 1760, 1, p. 48; 5, p. 261. Type, by tautonomy,
Limosa Brisson = Scolopax limosa Linnaeus.

Limosa lapponica baueri Naumann

Limosa baueri Naumann, Naturgeschichte Vogel Deutschlands, 8, 1836, p.
429. (New Holland.)

Pacific Godwit

Limosa lapponica novaezealandiae Gray in Rothschild, Avifauna of Laysan,
etc., p. 307; Henshaw, Birds Hawaiian Islands, etc., p. 93; Bryan, Key
Birds Hawaiian Group, p. 27; Perkins, Fauna Hawaiiensis (Aves), p. 45.

Range. Northeastern Asia and northwestern North America;
south in winter to islands of the Pacific and Australia.

Recorded in the Hawaiian Group (as an occasional visitor on migration)
from Laysan (Schauinsland), Kauai (W. A. Bryan), Maui (Ridgway).

Genus Heteroscelus — Tatlers

Heteroscelus Baird, Reports, Explorations and Surveys for a Railroad from the
Mississippi River to the Pacific Ocean, 9, 1858, pp. xxii, xlvii, 728, 734.
Type, by monotypy, Totanus brevipies Vieillot.

112 bulletin: museum of comparative zoology

Heteroscelus incanus (Gmelin)

Scolopax incana Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 658. (Eimeo

(= Moorea) and Palmerston Islands.)
Wandering Tattler, Ulili
Heteractitis incanus (Gmelin) in Rothschild, Avifauna of Laysan, etc., p. 255;

Henshaw, Birds Hawaiian Islands, etc., p. 92; Bryan, Key Birds Hawaiian

Group, p. 27; Perkins, Fauna Hawaiiensis (Aves), p. 450; Totanus incanus

in Wilson and Evans, Aves Hawaiiensis, p. 151.

Range. Alaska; south in winter to the west coast of America to
Ecuador and to the islands of the Pacific, sometimes to Australia.

Recorded in the Hawaiian Group (as a regular winter resident) from
Midway, Laysan, Lisianski, Necker, French Frigate Shoal, Niihau,
Kauai, Oahu, Maui, Hawaii," .... frequenting the rocky shores of all
islands" (Henshaw). Usually arrive in August and leave in May.
Though certain birds may remain all summer, there is no breeding
record for the islands.

Genus Arenaria — Turnstones

Arenaria Brisson, Omithologie, 1760, 1, p. 48; 5, p. 132. Type, by tautonomy,
Arenaria Brisson = Tringa interpret Linnaeus.

Arenaria interpres interpres (Linnaeus)

Tringa interpres Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 149. (Europe
and North America; type locality restricted to Gotland Island, Sweden,
by Hartert, Vogel Palaartischen Fauna, p. 1566.)

Turnstone, Akekeke

Tringa oahuensis Bloxam, Voyage "Blonde", 1826, p. 132.

Range. Northern Europe and northern Asia and Alaska west of
Point Barrow; south in winter to Africa, India, China and islands of
the Pacific.

Recorded in the Hawaiian Group (as a regular winter resident) from
Midway, Lisianski, Laysan, French Frigate Shoal, Necker, Niihau,
Kauai, Oahu, Molokai, Maui, Hawaii, "All the islands" (Perkins).
Usually arrive in August or September and depart in April or May.
Certain birds remain all summer, some in flocks at 6000 feet on Halea-
kala, Maui and in the crater of Kilauea, Hawaii. Subspecific identifica-
tion of winter birds is impossible. It seems more probable that most
of the birds to be seen in the Hawaiian Islands are interpres, though in-
dividuals of morinella, which is the bird that breeds in North America
east of Point Barrow, Alaska, may also reach the islands.

BRYAN AND GREENWAY: HAWAIIAN' BIRDS 113

Genus Capella — Snipe

Capella Frenzel, Beschreibung Vogel und Eyer . . . Wittenborg . . . , 1801,
p. 58. Type, by nionotypy, Scolopax coelestis Frenzel = Scolopax gallinago
Linnaeus.

( apella delicata (Ord)

Scolopax delicata Ord in Wilson, American Ornithology, 9, 1825, p. 218.

(Pennsylvania.)
Wilson's Snipe, Jack Snipe
Gallinago delicata in Rothschild, Avifauna of Laysan, etc., p. 253; Henshaw,

Birds Hawaiian Islands, p. 94; Perkins, Fauna Hawaiiensis (Aves), p. 451.

Range. Arctic and temperate North America; south in winter to
South America.

Recorded in the Hawaiian Group (as an occasional visitor on migration)
from Laysan, Oahu, Molokai, Maui, Hawaii.

Genus Crocethia — Sanderling

Crocethia Billberg, Synopsis Fauna Scandanaviae, 1, pt. 2, 1828, p. 132. Type
by nionotypy, Charadrius calidris Linnaeus = Trynga alba Pallas.

Crocethia alba (Pallas)

Trynga alba Pallas in Vroeg, Beredeneerde Catalogus, etc., Adumbratiunculae,
1764, N. 320, p. 7. ("Noordsche Zeekusten" = coast of northern Holland.)

Sanderling, Hunakai

Calidris aretiaria (Linnaeus) in Rothschild, Avifauna of Laysan, etc., p. 259;
Wilson and Evans, Aves Hawaiiensis, p. 153, 159; Henshaw, Birds
Hawaiian Islands, etc., p. 93; Perkins, Fauna Hawaiiensis (Aves), p. 451,
and other works.

Range. Arctic coasts of Europe, Asia, America ; south in winter to
temperate zone and southern hemisphere — Cape of Good Hope,
Malaya, Australia, Islands of the Pacific, southern South America.

Recorded in the Hawaiian Group (as a regular winter resident) from
Laysan, Niihau, Kauai, Maui (Ridgway), Oahu, Molokai, Hawaii,
"Most, and probably all, islands" (Perkins).

We have followed the A. O. U. Check-list and Peters (Birds of the
^Yorld) in recognizing but one form of this species. If more are recog-
nized the name of the bird which visits the islands will be Crocethia
alba tridactyla Pallas, the supposed form which breeds in northwestern
North America.

114 bulletin: museum of comparative zoology

Genus Erolia — Sandpipers

Erolia Vieillot, Analyse nouvelle ornithologie . . . 1816, p. 55. Type, by
monotypy, Erolia variegata Vieillot = Scolopax testacea Pallas.

Erolia acuminata (Horsfield)

Totanus acuminatum Horsfield, Transactions Linnaean Soc. London, 13, pt. 1,

1821, p. 192. (Java.)
Sharp-tailed Sandpiper
Heteropygia acuminata in Rothschild, Avifauna Laysan, etc., p. 255, 307;

Tringa acuminata in Henshaw, Birds Hawaiian Islands, p. 94; Perkins,

Fauna Hawaiiensis (Aves), p. 451.

Range. Eastern Asia; south in winter to south Pacific islands,
Malaya, New Guinea, Australia.

Recorded in the Hawaiian Group (as an occasional visitor on migra-
tion) from Midway, Laysan, Kauai, Oahu, Molokai (specimen in
Bishop Museum), Maui, Hawaii.

Erolia melanotos (Vieillot)

Tringa melanotos Vieillot, Nouveau Dictionnaire Hist. Nat., 34, 1819, p. 462.

(Paraguay.)
Pectoral Sandpiper
Tringa maculata in Henshaw, Birds Hawaiian Islands, etc., p. 94; Perkins,

Fauna Hawaiiensis (Aves), p. 451.

Range. Arctic zone of eastern Asia and North America; south in
winter to South America.

Recorded in the Hawaiian Group (as an accidental visitor on migration)
from Oahu (W. A. Bryan), Hawaii.

Genus Himantopus— Stilts

Himantopus Brisson, Ornithologie, 1760, 1, p. 46; 5, p. 33. Type, by tau-
tonomy, Himantopus Brisson = Charadrius himantopus Linnaeus.

Himantopus himantopus knudseni Stejneger

Himantopus knudseni Stejneger, Proceedings U. S. Nat. Mus., 10, 1887, p. 81,

pi. 6, fig. 2. (Kauai, Hawaiian Islands.)
Hawaiian Stilt, Aeo, Kukuluaeo

Range. Hawaiian Islands, where recorded from Niihau, Kauai,
Oahu, Molokai, Maui near lakes and ponds or on exposed mud flats.
"I have never seen it, however, upon Hawaii, nor have I been able to
learn of its presence there" (Henshaw).

B RY A N A ND GR E E N W A V : 1 1 A W AIIAN BIRDS 11 5

Family PHALAROPODIDAE

Genus Phalaropus — Phalaropes

Phalaropus Brisson, Ornithologie, 1760, 1, p. 50; 6, p. 12. Type, by tautonomy,
Phalaropus Brisson = Tringa fulicaria Linnaeus.

Phalaropus fulicarius (Linnaeus)

Tringa fulicaria Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 148. (Hudson

Bay ex Edwards.)
Red Phalarope
Crymophilus fulicarius in Rothschild, Avifauna Laysan, etc., p. 251; Henshaw,

Birds Hawaiian Islands, p. 95; Bryan, Key Birds Hawaiian Group, p. 25.

Range. Circumpolar in arctic; south in winter to seas off the coasts
of West Africa and Chile.

Recorded in the Hawaiian Group (as an irregular visitor on migration)
from Laysan, Kauai, Oahu, Maui, Hawaii, "evidently accompanied
the Akekeke (Arenaria interpres) to the uplands to feed" (!) (Hen-
shaw).

Phalaropus lobatus (Linnaeus)

Tringa tobata {Lobata in Emendanda, p. 824) Linnaeus, Systema Naturae, ed.

10, 1, 1758, p. 148. (Hudson Bay ex Edwards.)
Northern Phalarope

Range. Circumpolar in Arctic Zone; south in winter to oceans in
southern hemisphere.

Recorded in Hawaiian Group (as a chance visitor on migration) from
Kauai.

Family LARIDAE

Genus Larus — Gulls

Larus Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 136. Type, by subsequent
designation, Larus marinus Linnaeus (Selby, Catalogue generic and sub-
generic types . . . Aves, 1840, p. 48).

Larus delawarensis Ord

Larus delawarensis Ord in Guthrie's New Geographical . . . Grammar, etc.,
2nd American ed., 2, 1815, p. 319. (Delaware River, below Philadelphia.)
Ring-billed Gull

116 bulletin: museum of comparative zoology

Range. Northwestern and northeastern North America; south in
winter, sometimes as far as the Greater Antilles and Mexico.

Recorded in the Hawaiian Group (as an accidental visitor) from
Molokai and Maui or Hawaii (Bryan).

Larus argentatus smithsonianus Coues

Larus smithsonianus Coues, Proceedings Academy Nat. Sci., Philadelphia,

1862, p. 296. (Eastern and western coasts of North America.)
Herring Gull

Range. North America in arctic and northern temperate zones;
south in winter, sometimes to northern tropics.

Recorded in Hawaiian Group (as an accidental visitor) from Laysan.

Larus californicus Lawrence

Larus californicus Lawrence, Annals Lyceum Nat. Hist., New York, 6, 1854,

p. 79. (Near Stockton, California.)
California Gidl

Range. Northwestern North America from the Mackenzie River
east to Saskatchewan, south to west central California and north-
western Wyoming; south in winter to southern California and western
Mexico.

Recorded in the Hawaiian Group (as an accidental visitor) from Maui
or Hawaii (Bryan, 1901).

Larus pipixcan Wagler

Larus pipixcan Wagler, Isis von Oken, 1831, col. 515. (Mexico.)
Franklin's Gull

Larus franklini Swainson and Richardson in Henshaw, Birds of Hawaiian
Islands, p. 127; Bryan, Key Birds Hawaiian Group, p. 6.

Range. Interior of northwestern North America; south in winter
to coasts of the Gulf of Mexico and western coasts of South America.

Recorded in the Hawaiian Group (as an accidental visitor) from Maui
(one record).

Larus glaucescens Naumann

Lams glaucescens Naumann, Naturgeschichte Vogel Deutschlands, etc., 10,

1840, p. 351. ("Nord-Amerika".)
Glaucous-uinged Gull

BRYAX AM) GREENWAT: HAWAIIAN BIRDS 117

Range. Northeastern Asia arid northwestern North America;
south in winter to China and Japan and the Aleutian Islands to Cali-
fornia.

Recorded in the Hawaiian Group (as an accidental visitor) from Lay-
san, Hawaii, "Probably . . . other islands, especially to Oahu" (Hen-
shaw).

This gull sometimes follows ships ta the Hawaiian Islands.

Larus hyperborevs Gunnerus

Larus hyperboreus Gunnerus, in Leem . . . Beskrivelse Finmarkens Lapper,

etc., 1767, p. 226, (footnote). (Northern Norway.)
Glaucous Gull, Point Barrow Gull

Larus glaucus Brunnich, Ornithologia borealis, etc., 1764, p. 44.
Larus barrorianus Ridgway, Auk, 3, 1886, p. 330; in Henshaw, Birds Hawaiian

Islands, p. 126; Bryan, Key Birds Hawaiian Group, p. 6.

Range. Circumpolar in the Arctic; south in winter to coast of
temperate zones, straying sometimes far.

Recorded in Hawaiian Group (as an accidental visitor) from Laysan,
Kauai, Lanai, Maui.

Larus Philadelphia (Ord)

Sterna Philadelphia Ord in Guthrie's New Geographical . . . Grammar, etc.,
2nd American ed., 2, 1815, p. 319. (No locality = near Philadelphia,
Pennsylvania.)

Bonaparte s Gull

Range. Northwestern North America ; winters on the Atlantic and
Pacific coasts.

Recorded in the Hawaiian Group (as an accidental visitor) from Kauai
(Rothschild). '

Genus Rissa — Kitti wakes

Rissa Stephens, in Shaw's General Zoology, 13, pt. 1, 1826, p. 180. Type, by
monotypy, Rissa brunnichii Stephens = Larus tridactylus Linnaeus.

Rissa tridactyla pollicaris Ridgway

Rissa tridactyla pollicaris attributed to Stejneger by Ridgway in Baird,
Brewer and Ridgway, Water Birds North America, 2, 1884, p. 202.
(Kotzebue Sound, Alaska.)

Western Kittiwake

118 bulletin: museum of comparative zoology

Range. Eastern Siberia, Islands of Behring Sea, Alaska; south in
winter to Japan and California.

Recorded in the Hawaiian Group (as an accidental visitor) from
Laysan (fragments in Bishop Museum).

Genus Sterna — Terns

Sterna Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 137. Type, by tau-
tonomy, Sterna hirundo Linnaeus.

Sterna paradisaea Pontoppidan

Sterna Paradisaea Pontoppidan, Danske Atlas, 1, 1763, p. 622. (Christansoe,

Denmark ex Briinnich 1764.)
Arctic Tern

Range. Circumpolar in Arctic and parts of northern temperate
zones; south in winter to the Antarctic Ocean.

Recorded in the Hawaiian Group ( as an accidental visitor) from
Oahu, Hawaii (Henshaw).

Sterna sumatrana sumatrana Raffles

Sterna sumatrana Raffles, Transactions Linnaean Soc. London, 13, pt. 2, 1822,

p. 329. (Sumatra.)
Black naped Tern
Sterna melanauchen Temminck in W. A. Bryan, Birds Hawaiian Group, p. 8;

Perkins, Fauna Hawaiiensis (Aves), p. 464; Henshaw, Birds Hawaiian

Islands, p. 124.

Range. Western and southern Pacific and eastern Indian Oceans.
Recorded in the Hawaiian Group (as an accidental visitor) from
Kauai, Hawaii.

Sterna anaethetus lunata Peale

Sterna lunata Peale, U. S. Exploring Expedition, 8, 1848, p. 277. (Vincennes

Island = Kauehi Island, Tuamotus.)
Bridled Tern, Gray backed Tern, Pakalakala
Haliplana lunata in Rothschild, Avifauna of Laysan, etc., v, vii, ix, 37.

Range. Pacific Ocean from the Hawaiian Islands south to the
Moluccas.

Recorded in the Hawaiian Group from Laysan (breeding), ? Lisianski,
Gardner (sight, Rothschild), ? French Frigate Shoal (Fisher ? ex-
Rothschild), Necker, Niihau, Kauai, Oahu, Hawaii.

BRYAN AND GREEXWAY : HAWAIIAN BIRDS ] 1!)

Sterna fuscata oahuensis Bloxam

Sterna Oahiu nsis Bloxam, Voyage "Blonde", 1826, p. 251. (Oahu.)

Wideawake, Sooty Tern, Ewa< wa

Haliplana fuMginosa. (Gmelin) in Rothschild, Avifauna of Laysan, etc., vii,
viii, 39; Sterna fuliginosa (Gmelin) in Wilson andEvans, AvesHawaiiensis-
p. 137; Bryan, Key Birds Hawaiian Group, p. 8; Perkins, Fauna Hawai-
iensis (Aves), p. 464; Henshaw, Birds Hawaiian Islands, p. 122, et al.

Range. Pacific Ocean from the Bonin Islands and the Hawaiian
Islands southward.

Recorded in the Hawaiian Group from Midway (breeding), Pearl and
Hermes Reef, Lisianski, Laysan (breeding), ? Gardner (sight, Roths-
child), French Frigate Shoal (breeding), Necker (breeding), Bird Rock
(Nihoa), Kaula, Kauai, Oahu (Moku Manu).

Genus Procelsterna — Terns

Procelsterna Lafresnaye, Magazin Zoologie, 1842, Oiseaux, pi. 29, p. 1. Type,
by monotypy, Procelsterna tereticollis Lafresnaye = Sterna teretirostris
Lafresnaye.

Procelsterna certjlea saxatilis W. K. Fisher

Procelsterna saxatilis W. K. Fisher, Proceedings U. S. Nat. Mus., 26, 1903,

p. 559. (Necker Island.)
Necker Island Tern

Range. Marcus Island and western Hawaiian Islands.
Recorded in the Hawaiian Group from French Frigate Shoal, Necker
(breeding), Bird Island (Nihoa), Kaula.

Genus Ano us Stephens — Noddies

Anoiis Stephens, in Shaw's General Zoology, 13, pt. 1, 1826, p. 139. Type, by
subsequent designation, Anoiis niger Stephens = Sterna stolida Linnaeus
(Gray, List Genera Birds, 1840, p. 79).

Anous stolidus pileatus (Scopoli)

Sterna pileata Scopoli, Deliciae florae et faunae insubricae, etc., fascic. 2,
1786, p. 92. (No type locality — Philippine Islands ex Sonnerat, Voyage
aux Indes, etc., 1782.)

Noddy

Anous stolidus (Linnaeus) in Rothschild, Avifauna of Laysan, etc., p. 41 ; Wilson
and Evans, Aves Hawaiiensis, p. 141 ; Bryan, Key Birds Hawaiian Group,
p. 9; Perkins, Fauna Hawaiiensis (Aves), p. 464; Henshaw, Birds Ha-
waiian Islands, p. 124.

120 bulletin: museum of comparative zoology

Range. Indian and western Pacific Oceans, Seychelles and Mada-
gascar, Bonin and Hawaiian Islands south to Australia.

Recorded in the Hawaiian Group from Midway (breeding), Laysan
(breeding), French Frigate Shoal, Necker, Bird Island (Nihoa),
Kaula Lehua (breeding), Oahu.

Anous minutus melanogenys G. R. Gray

Anous melanogenys G. R. Gray, Genera Birds, 3, 1846, pi. 182. (No type
locality; figure agrees with Hawaiian bird teste Hartert.)

Hawaiian Tern, Noio

Anous hawaiiensis Rothschild in Rothschild, Avifauna of Laysan, etc., vii, xi,
43, 285; Wilson and Evans, Aves Hawaiiensis, p. 143; Perkins, Fauna
Hawaiiensis (Aves), p. 464; Henshaw, Birds Hawaiian Islands, p. 125;
Microanous hawaiiensis Bryan, Key Birds Hawaiian Group, p. 9; et al.

Range. Hawaiian Islands.

Recorded in the Hawaiian Group from Midway (breeding), Laysan
(breeding), Lisianski (breeding), Necker, Bird Rock (Nihoa), Lehua,
Niihau (sight, Rothschild), Kauai, Oahu (Mokuloa and Kaohi Kaipu)
? Molokai (W. K. Fisher), Hawaii.

Genus Gygis — Fairy Terns

Gygis Wagler, Isis von Oken, 1832, col. 1223. Type, by monotypy, Sterna
Candida Gmelin.

Gygis alba rothschildi Hartert

Gygis alba rothschildi Hartert, Novitates Zoologicae, 34, 1927, p. 18. (Laysan
Island.)

White Tern, Love Bird

Gygis alba (Sparrman) or Gygis Candida Wagler in Rothschild, Avifauna of
Laysan, etc., pp. vii, xiii, xiv, 35, 285; Wilson and Evans, Aves Ha-
waiiensis, p. 145; Perkins, Fauna Hawaiiensis, p. 145; Henshaw, Birds
Hawaiian Group, p. 126; Gygis alba kitllitzi Hartert in Bryan, Key Birds
Hawaiian Group, p. 9; et al.

Range. Islands northwest of Kauai.

Recorded in the Hawaiian Group from Midway (breeding), Laysan
(breeding), Lisianski, French Frigate Shoal, Necker (breeding),
Bird Rock (Nihoa), Kaula.

BRYAN AND GREENWAY: HAWAIIAN UIRDS 121

Order STRIGIFORMES

Family STRIGIDAE

Genus Asio Brisson

Asio Brisson, Ornithologie, 1, 1760, p. 28. Type, by tautonomy, Asio Brisson
= Strix otus Linnaeus.

Asio flammeus sandwichensis (Bloxam)

Strix sandwnchensis Bloxam, Voyage "Blonde", 1826, p. 250. (Hawaiian

Islands.)
Hawaiian Owl, Pueo
Asio acciplrinus in Wilson and Evans, Aves Hawaiiensis, p. 133; Perkins,

Fauna Hawaiiensis (Aves), p. 448.

Range. Kauai (breeding), Oahu (breeding), Molokai (? breeding),
Maui, Hawaii (breeding).

Order CORACIIFORMES

Family ALCEDINIDAE

Genus Ceryle — Kingfishers

Ceryle Boie, Isis von Oken, 1828, col. 316. Type, by subsequent designation ,
Alcedo rudis Linnaeus, Gray, List Genera Birds, 1840, p. 11.

Ceryle alcyon caurina Grinnell

Ceryle alcyon caurina Grinnell, University California Publications, Zool., 6>
1910, p. 388. (Graveyard Point, Montague Island, Prince William
Sound, Alaska.)

Western Belted Kingfisher

Ceryle alcyon (Linnaeus) in Henshaw, Birds Hawaiian Group, p. 77.

Range. Alaska south to southern California; south to northern
and central Mexico in winter.

Recorded in the Hawaiian Group (as an accidental visitor) from
Hawaii.

Order PASSERIFORMES— Perching Birds

Family CORVIDAE

Genus Corvus — Crows

Corvus Linnaeus, Systema Naturae, ed. 10, 1, 1758, p. 105. Type, by Linnaean
tantonymy, Corvus corax Linnaeus.

122 bulletin: museum of comparative zoology

Corvus hawaiiensis Peale

Corvus hawaiiensis Peale, U. S. Exploring Expedition, 8, 1848, p. 106. (Kara-
kakua [Kealakekua] Bay, Hawaii.)

Hawaiian Crow, Alala

Corvus tropicus "Linnaeua", Bloxam in Voyage "Bonde", 1826, p. 250 (Sand-
wich Islands) is a nomen nudum. Corvus tropicus Gmelin to which Bloxam
refers is not identifiable.

Corvus tropicus Gm. in Wilson and Evans, Aves Hawaiiensis, p. 1 ; Henshaw,
Birds Hawaiian Possessions, etc., p. 36.

Range. Edges of mountain forest from 1000 to 6000 feet in Kona
and Kau districts of the Island of Hawaii.

Family MUSCICAPIDAE— Flycatchers, Thrushes, Warblers
Subfamily TURDINAE

Genus Phaeornis — Hawaiian Thrushes

Phaeornis Sclater, Ibis, 1, 1859, p. 327. Type, by original designation and
monotypy, Phaeornis obscura (Gmelin).

Phaeornis obscura obscura (Gmelin)

Muscicapa obscura Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 945. "(in

insulis Sandwich.")
Ornao, Oman, Hawaii Thrush, Kamao, A-Maui.

Range. Endemic to Island of Hawaii.

Phaeornis obscura lanaiensis Wilson

Phaeornis lanaiensis Wilson, Annals and Magazine Nat. Hist. (6), 7, 1891 »

p. 460. (Lanai.)
Lanai Thrush, OXomao, Olomau

Range. Mountain forests of Lanai. Now very rare.

Phaeornis obscura rutha W. A. Bryan

Phaeornis rutha W. A. Bryan, Occasional Papers B. P. Bishop Mus., Honolulu,

4, no. 2, 1908, p. 81. (Molokai.)
Molokai Thrush, Olomao

Range. Mountain forests of Molokai.

This poorly marked form averages somewhat larger (wing averages
92 mm. and 95 mm.) and slightly darker than lanaiensis.

BRYAN AND GRKKNWAV: HAWAIIAN BIRDS 123

Phaeornis ? obscurus oahensis Wilson and Evans

Phaeornis oahensis Wilson and Evans, Aves Hawaiiensis (Introduction),

1899, p. XIII (Oahu).
Oahu Thru ah

Range. Formerly mountain forests of Oahu. Now extinct.

Specimens are supposed to have been collected by Andrew Bloxam,
naturalist on H. M. S. 'Blonde', but they have disappeared. Wilson
and Evans described the bird from Bloxam's manuscript notes.

Phaeornis obscura myadestina Stejneger

Phaeornis myadestina Stejneger, Proceed. U. S. Nat. Mus., 10, 1887 (1888),
p. 90. (Kauai.)

Phaeornis myiadestina Wilson and Evans, Aves Hawaiiensis, p. 117; Roths-
child, Avifauna Laysan, p. 63.

Kauai Thrush, Kamau, Kamao

Range. Forests of Kauai.

Phaeornis palmeri Rothschild

Phaeornis palmeri Rothschild, Avifauna Laysan Island, etc., pt. 2, 1893, p. 67.

(Halemanu, Kauai.)
Puaiohi

Range. Region of Halemanu, Kauai. Never recorded from other
parts of the island. Now very rare and possibly extinct.

This bird, because of its voice and habits, which differ from those
of other members of the genus (fide Perkins et al.), and because it
apparently breeds in the same region as myadestina, we must treat
as a distinct species.

Subfamily SYLVIINAE

Genus Acrocephalus- — Miller Birds

Acrocephalus Naumann, Land u. Wasservogel nordl. Deutschland, Nachtrage,
4, 1811, p. 199. Type, by subsequent designation, Sylvia turdoides Meyer
= Acrocephalus arundinaceus (Linnaeus).

We can see no reason for retaining the genus Conopoderas Billberg.
Some of the island forms are closer to the type of Acrocephalus than
to that of Conopoderas.

124 bulletin: museum of comparative zoology

ACROCEPHALUS FAMILIARIS (Rothschild)

Tatare familiaris Rothschild, Annals and Magazine Nat. Hist. (6), 10, 1892,

p. 109. (Laysan Island.)
Laysan Miller Bird

Range. Formerly Laysan Island, now extinct.

Acrocephalus kingi ( Wetmore)

Conopoderas kingi Wetmore, Condor, 26, 1924, p. 177. (Nihoa.)
Nihoa Miller Bird

Range. Nihoa (Bird Rock).

In employing binomials we do not mean to imply that these forms
are distinct species. The present application of specific names is
unsatisfactory. This genus needs revision.

Subfamily MUSCICAPINAE

Genus Chasiempis Cabanis

Chasiempis Cabanis, Archiv fur Naturgeschiehte, 13, 1847, Bd. 1, p. 207.
Type, by original designation, "Muscicapa sandvicensis Latham" =
Chasiempis sandwichensis (Gmelin).

Though Cabanis cited Muscicapa sandvicensis Latham, Latham
himself never used this name. Gmelin 's description of Muscicapa
sandwichensis was taken from Latham's "Sandwich Fly-catcher."

Chasiempis sandwichensis sclateri Ridgway

Chasiempis sclateri Ridgway, Proceed. U. S. Nat. Mus., 4, 1881, (1882), p. 337.

(Waimea, Kauai.)
Kauai Elepaio, Apekepeke, Amakahi, Kahuna-Ka-lai-woa
Chasiempis dolei Stejneger, Proceed. U. S. Nat. Mus., 10, 1887 (1888), p. 90.

(Kauai.)

Range. Forests of Kauai.

Both Hawaiians and scientists were long under the impression that
the white-rumped form (i\makahi or Elepaio) and the brown-rumped
form (Apekepeke) were distinct species. Perkins showed that the
latter is an immature plumage in which the birds sometimes breed.

BRYAN AND GREENWAY: HAWAIIAN BIRDS 125

Chasiempis sandwichensis gayi Wilson

Chasiempis gayi Wilson, Proceed. Zool. Soc. London, 1891, p. 165. (Oahu.)
Oahu Elepaio

Range. Forests of Oahu.

Chasiempis sandwichensis sandwichensis (Gmelin)

Muscicapa sandwichensis Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 945.

("in insulis Sandwich" = Hawaii.)
Hawaii Elepaio, Ono-Ka-Ia
Chasiempis rirfgwayi Stejneger, Proceed. U. S. Nat. Mus., 10, 1887 (1888),

p. 87, in key, p. 89. (Hawaii.)
Chasiempis ibidis Stejneger, Proceed. U. S. Nat. Mus., 10, 1887 (1888), p. 87,

in key, p. 89.

Range. Forests of Hawaii.

Stejneger's Chasiempis ibidis refers to plate 1, figure 2 in the Ibis
of 1S85. This specimen had no locality other than "Chili," as Sclater
remarks.

Subfamily DREPANIDIXAE

Genus Viridonia

Yiridonia Rothschild, Annals and Magazine Nat. Hist. (6), 10, 1892, p. 112.
Type, by monotypy, Viridonia sagittirostris Rothschild.

Viridonia sagittirostris Rothschild

Viridonia sagittirostris Rothschild, Annals and Magazine Nat. Hist. (6), 10,

1892, p. 112. (Lower Hilo slope of Mauna Kea, Hawaii.)
Green Solitaire

Range. Mountain rain forests of the windward side of Hawaii near
the Wailuku River at about 1200 to 4000 feet.

Genus Palmeria

Palmeria Rothschild, Ibis, 1893, p. 113. Type, by original designation and
monotypy, Palmeria mirabilis Rothschild = P. dolei I Wilson).

126 bulletin: museum of comparative zoology

Palmeria dolei (Wilson)

Himatione dolei Wilson, Proceed. Zool. Soc. London, 1891, p. 166. (Maui.)

Akohekohe, Crested honey eater

Palmeria mirabilis Rothschild, Ibis, 1893, p. 113.

Range. Mountain forests of Molokai and Maui. Now very rare.

Genus Himatione

Himatione Cabanis, Museum Heineanum, Th. 1, 1850, p. 99. Type, by subse-
quent designation, Himatione sanguinea (Gmelin).

Himatione sanguinea fraithii Rothschild

Himatione Fraithii Rothschild, Annals and Magazine Nat. Hist. (6), 10, 1892,

p. 109. (Laysan Island.)
Laysan Honey Creeper

Range. Formerly Laysan Island, now extinct.

Himatione sanguinea sanguinea (Gmelin)

Certhia sanguinea Gmelin, Systema Naturae, 1, pt. 1, 1788, p. 479. (Sandwich

Islands.)
Apapane, Akakani

Range. Mountain forests of ?Niihau, Kauai, Oahu, Molokai,
Maui, Hawaii.

Genus Vestiaria

Vestiaria Fleming, Philosophy of Zoology, 2, 1822, p. 246. Type, by original
designation, Certhia vestiaria = Vestiaria coccinea (Forster).

Vestiaria coccinea coccinea (Forster)

Certhia coccinea Forster, Goettingisches Magazin Wissenschaften, etc., 1,

1780, p. 347.
Iiwi, Iiwi popolo, Iiwi polena (for various phases of plumage).
Vestiaria coccinea suavis Bangs, Proceed. Biol. Soc. Wash., 24, 1911, p. 29.

(Molokai.)

Range. Mountain forests of Kauai, Oahu, Molokai, Maui, Lanai,
Hawaii.

BRYAN AND GREENWAY: HAWAIIAN BIRDS 127

Tor full synonymy see Rothschild, Avifauna Laysan Island, etc.

Vestiaria coccinea suavis Bangs was described as paler, with a larger
bill. In our opinion, the former is an individual variation and the latter
is non-existent. In our short series (28) there is an average difference
of 2 mm. in wing length, but this is not sufficient for formal recognition.

Genus Drepaxis — Hawaiian Honey Creepers

Drepanis Temminck, Manual d'Ornithologie, ed. 2, 1, 1820, p. LXXXVI .
Type, by subsequent designation (Gray, 1840), Certhia pacifica Gmelin.

Drepaxis pacifica (Gmelin)

Certhia pacifica Gm., Systema Naturae, 1, pt. 1, 1788, p. 470. ("in insulis

amicis" in error ex Latham. Hawaiian Islands.)
Mamo, Hoha or Hoho (so rendered by older writers).

Range. Formerly mountain forests of Hawaii (old records for
Kauai are almost certainly in error). Now extinct; last specimen seen
at Kaumana 1000-1500 feet above Hilo, 1898.

Drepaxis fuxerea Newton

Drepanis funerea Newton, Proceed. Zool. Soc. London, p. 690. (Molokai.)

Ma?no, Oo-nuku-umu, Hoa, Black Mamo, Perkins' Mamo

Drepanorhamphus funerea Newton in Rothschild, Avifauna Laysan Island,

etc., p. 165; Bryan, Birds Hawaiian Group, p. 42; Perkins, Fauna Ha-

waiiensis (Avesj, p. 402.

Range. Mountain forests of Molokai.

The genus Drepanorhamphus was established by Rothschild (Avi-
fauna Laysan, etc., pt. 3, 1900, p. 163. Type, by monotypy, Drepanis
funerea Newton). The characters that separate it from Drepanis are
the lack of the long, decomposed, yellow under tail coverts and the
shape of the nostril which is long and narrow, not rounded. It seems
to us that they are alike in so many ways that they form a single genus
separating them from other genera, but that they were probably dis-
tinct species.

Genus Hemigxathus — Akialoa

Hemignathus Lichtenstein, Abhandlungen Konig. Akademie Wissensch
Berlin, 1839, p. 449. Type, by subsequent designation, Hemignathus
lucidus Licht. (Gray, List Genera Birds, 1841, p. 16).

128 bulletin: museum of comparative zoology

In our opinion, the group with long lower mandibles and those with
short lower mandibles may well be considered as congeneric. If it is
desired to separate them generically then the former will require a
new generic name and the latter (heretofore known as Heterorhynchus
will have to be called Hemignathus, since the two groups, as named
heretofore, have the same type as designated by Gray and Lafresnaye.
Rothschild's arguments (Avif. Laysan Id., etc., p. 79) have no force
under the rules of zoological nomenclature (Art. 30, II, g).

Hemignathus obscurus procerus Cabanis

Hemignathus procerus Cabanis, Journal f. Ornith., 1889, p. 331. (Kauai.)
Kauai Akialoa, "Iiwi"

Range. Mountain forests of Kauai from the lowest forest zone
600-900 feet to 4000 feet.

Hemignathus obscurus ellisianus (Gray)

Drepanis {Hemignathus) ellisiana Gray, Cat. Birds Tropical Islands Pacific,

1860, p. 9. (Oahu.)
Oa.hu Akialoa, Iiwi (Jibi or Kipi)
Hemignathus lichtensteini Wilson in Aves Hawaiiensis, p. 65, and Perkins,

Fauna Hawaiiensis (Aves), p. 425.

Range. Formerly mountain forests of Oahu; now extremely rare;
perhaps confined to Waianae Mountains (Northwood).

Hemignathus obscurus lanaiensis Rothschild

Hemignathus lanaiensis Rothschild, Bull. Brit. Ornith. Club, 1, 1893, p. xxiv.

(Lanai.)
Lanai Akialoa

Range. Formerly mountain forests of Lanai, now very rare, pos-
sibly extinct.

Hemignathus obscurus obscurus (Gmelin)

Certhia obscura Gm., Systema Naturae, 1, pt, 1, 1788, p. 470. (Sandwich

Islands = Hawaii.)
Haivaii Akialoa

Range. Formerly mountain forests of Hawaii, now probably con-
fined to localized areas in the rain forests of the windward side (see
Baldwin, 1941, p. 19).

BRYAN AND (iKKF.NW AY: HAWAIIAN BIRDS 129

Hemignathus lucidus iiaxapepe Wilson

Hemignathus hanapepe Wilson, Annals Magazine Nat. Hist. (6), 4, 1889,

p. 401. (Kauai.)
Nukupu'u
Heterorhgnrhus hanapt /« in Rothschild, Avifauna Laysan, etc., p. 101 ; Perkins,

Fauna Hawaiiensis (Aves), p. 430; Henshaw, Birds Hawaiian Islands,

p. 43.

Range. Mountain forests of Kauai in Waimea district near the
Hanapepe River, 2000 to 3000 feet, very rare and local.

Hemignathus lucidus lucidus Lichtenstein

Hemignathus lucidus Lichtenstein, Abhandlungen der Konigl. Academie
Wissensch. Berlin, 1S39, p. 451, pi. 5, figs. 2, 3. (Oahu.)

Oahu Akiapolaau

HeterorhyncJms lucidus in Rothschild Avifauna Laysan, etc., p. 105; Perkins,
Fauna Hawaiiensis (Aves), p. 430; Henshaw, Birds Hawaiian Islands,
p. 41.

Akiapolaau

Range. Formerly mountain forests of Oahu ; now almost certainly
extinct.

Ten specimens were thought to exist in museums but there are not
that many, for some have been misidentified.

Hemignathus lucidus affinis Rothschild

Hemignathus affinis Rothschild, Ibis, 1893, p. 112. (Maui.)
Maui Akiapolaau

Range. Recorded from forested slopes of Haleakala Mountain,
Maui, 4000 to 4500 feet. Now local and probably very rare.

Hemignathus lucidus wilsoni (Rothschild)

Heterorhynchus wilsoni Roths., Avifauna Laysan, etc., 1893, pt. II, p. 97.

(Hawaii).
Akiapolaau
Hemignathus olivaceus Lafr. in Wilson and Evans, Aves Hawaiiensis, p. 75.

Range. Mountain forests of Hawaii from about 1500 to 6700 feet.
Still exists but not common.

130 bulletin: museum of comparative zoology

Genus Chlorodrepanis — Amakihi

Chlorodrepanis Perkins in Wilson and Evans, Aves Hawaiiensis, pt. 7, p. xxi
(introduction), June 1899. Type, by subsequent designation, Chlorodre-
panis stejriegeri Wilson (Richmond, Proceed. U. S. Nat. Mus., 24, 1902,
p. 673).

Chlorodrepanis parva (Stejneger)

Himatione parva Stejneger, Proceedings U. S. Nat. Mus., 10, 1887, p. 94.

(Kauai.)
Alawi, Anauanii, Anianau
Oreomyza parva in Rothschild, Avifauna Laysan, etc., p. 119.

Range. Forests of Kauai.

This is a very curious little bird for which Wilson and Evans pro-
posed the name Rothschildia (pre-occupied). Mathews in 1925 re-
named it Magumma. Our treatment is based on that of Perkins
(Ibis, 1895 and Fauna Hawaiiensis (Aves), p. 411).

Chlorodrepanis virens stejnegeri (Wilson)

Himatione stejnegeri Wilson, Proceedings Zool. Soc. London, 1889 (1890),

p. 446. (Kauai.)
Kauai Amakihi

Range. Forests of Kauai.

Chlorodrepanis virens wilsoni (Rothschild)

Himatione wilsoni Rothsch., Bull. Brit. Ornith. Club, 1, 1893, p. xliii. (Maui.)

Molokai Amakihi, Lanai Amakihi, Maui Amakihi

Himatione kalaana Wilson and Evans, Aves Hawaiiensis, 1896, p. 28. (Molo-
kai.)

Himatione chloridoides Wilson and Evans, Aves Hawaiiensis, 1896, p. 28.
(Lanai.)

Range. Forests of Molokai, Lanai, Maui.

This form is very close to chloris but differs in having a slightly
larger bill and (in mature specimens) it has a slightly paler back.
Differences between specimens from Molokai, Lanai and Maui alleged
to exist (an ill-defined or well-defined yellow stripe from bill to eye as
well as the shade of green on the back) fall within the range of indi-
vidual variation.

BRYAN AND GREENWAT: HAWAIIAN BIRDS 131

Chlorodrepanis virens virens (Gmelin)

Certhia virens Gm., S}'stema Naturae, 1, pt. 1, 17S8, p. 479. (in insulis Sand-
wich = Hawaii.)
Hawaii Amakihi

Range. Forests of Hawaii.

Genus Loxops — Akakane

Loxops Cabanis, Archiv fur Naturgeschichte, Berlin, 13, 1847, Bd. 1, p. 330.
Type, by original' designation, Fringilla coccinea Gm. For synonymy see
Rothschild, Avifauna Laysan, etc., p. 167.

Loxops caeruleirostris (Wilson)

Chrysomitridops caeruleirostris Wilson, Proceedings Zool. Soc. London, 1889

(1890), p. 445. (Kauai.)
Ou-holoicai, Akekee (fide Perkins)
Chrysomitridops caeruleirostris in Wilson and Evans, Aves Hawaiiensis, p. 59.

Range. Forests of Kauai.

Loxops coccinea rufa (Bloxam)

Fringilla rufa Bloxam in Voyage "Blonde", 1826, p. 250. (Oahu.)
Akepa, Akepeuie

"Loxops wolstenkolmei" Roths. — the plate so lettered in Rothschild Avifauna
Laysan, etc., p. 177.

Range. Formerly forests of Oahu, now very rare and perhaps
extinct. Last record May 20, 1S93.

Loxops coccinea ochracea Rothschild

Loxops ochracea Rothschild, Ibis, 1893, p. 112. (Maui.)
Ochraceus or Maui Akt peuie

Range. Forests of Maui.

Loxops coccinea coccinea (Gmelin)

Fringilla coccinea Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 921. (in insulis

Sandwich = Hawaii.)
Akepa, Akepeuie

Range. Forests of Hawaii. "Now rare" (Baldwin).

132 bulletin: museum of comparative zoology

Genus Paroreomyza

Paroreomyza Perkins, Ibis, 1901, p. 583. Type, by original designation,
Himatione maculata Cabanis.

In our opinion, the forms formerly listed under this name and those
listed under the name Oreomystis Stejneger 1903 are congeneric. It is
true that females of the former lack the color of the males, but the
birds are otherwise alike from a generic point of view.

Paroreomyza bairdi bairdi (Stejneger)

Oreomyza bairdi Stejneger, Proceedings U. S. Nat. Mus., 10, 1887 (1888), p. 99.

(Kauai.)
Akikihi
Oreomyza bairdi in all works on Hawaiian birds.

Range. Forests of Kauai above 1000 feet.

Paroreomyza bairdi mana (Wilson)

Himatione mana Wilson, Annals and Magazine Nat. Hist. (6), 7, 1891, p. 460.

(Hawaii.)
Hawaii Creeper, Olive-green Creeper
Oreomyza mana in all works on Hawaiian birds.

Range. Forests of Hawaii except (fide Perkins) in lower elevations
on the Kona or leeward coast. Now uncommon (fide Baldwin 1941).

In our opinion "Oreomyza perkinsi" Rothschild is in all probability
a hybrid, Chlorodrepanis virens x Paroreomyza mana. It has been listed
with other hypothetical forms and records.

Paroreomyza maculata maculata (Cabanis)

Himatione maculata Cabanis, Museum Heineanum, Th. 1, 1850, p. 100, in

footnote. (Oahu.)
Oahu Creeper
Himatione maculata Cabanis in Wilson and Evans, Aves Hawaiensis, p. 43;

Oreomyza maculata in Rothschild, Avifauna Laysan Island, etc., p. 113;

W. A. Bryan, Key Birds Hawaiian Group, p. 48; Perkins, Fauna Ha-

waiiensis, (Aves) p. 417; Henshaw, Birds Hawaiian Islands, p. 50.

Range. Mountain forests of Oahu above 1500 feet. "Scarcest of
the five native perching birds which are likely to be seen." (North-
wood 1940).

BRYAN AND OKEENWAY : HAWAIIAN BIRDS 133

Paroreomyza maculata flammea (Wilson)

Loxops flarnmea Wilson, Proceedings Zool. Soc. London, 1889 (1890), p. 445.
(Kalae, Molokai.)

Kakawahie

Loxops flammea Wilson in Wilson and Evans, Aves Hawaiiensis, p. 39; Oreomyza
flammea Wilson in Rothschild, Avifauna Laysan Island, etc., p. 121;
W. A. Bryan, Key Birds Hawaiian Group, p. 48; Perkins, Fauna Hawaii-
ensis (Aves), p. 417; Henshaw, Birds Hawaiian Islands, p. 49.

Range. Mountain forests of Molokai above 1500 feet.

Paroreomyza maculata Montana (Wilson)

Himatione montana Wilson, Proceedings Zool. Soc. London, 1889 (1890), p.
446. (Lanai.)

Alauhiio

Himatione montana Wilson in Wilson and Evans, Aves Hawaiiensis, p. 45;
Oreomyza montana in Rothschild, Avifauna Laysan Island, etc., p. 117;
W. A. Bryan, Key Birds Hawaiian Group, p. 47; Perkins, Fauna Hawaii-
ensis (Aves), p. 417; Henshaw, Birds Hawaiian Islands, p. 51.

Range. Mountain forests of Lanai.

Paroreomyza maculata newtoni (Rothschild)

Himatione newtoni Rothschild, Bull. British Ornith. Club, 1, 1893, p. xlii.

(Maui.)
Maui Creeper
Himatione newtoni Rothschild in Wilson and Evans, Aves Hawaiiensis, p. 11;

Oreomyza newtoni Rothschild in Rothschild, Avifauna Laysan Island, etc.

p. 115; W. A. Bryan, Key Birds Hawaiian Group, p. 48; Perkins, Fauna

Hawaiiensis (Aves), p. 417; Henshaw, Birds Hawaiian Islands, p. 49.

Range. Mountain forests of Maui.

Genus Ciridops

Ciridops A. Newton, Nature, 45, 1892, p. 469. Type, by monotypy, Fringilla
anna Dole.

Ciridops anna (Dole)

Fringilla anna Dole, Hawaiian Almanac, 1879, p. 49. (Hawaii.)
Waaihaiuane

134 bulletin: museum of comparative zoology

Range. Formerly mountain forests of Hawaii. Recorded from
Kona and Hilo districts in or near Loulu or Hawane palms (Prit-
chardia). Never common within the memory of man, it is now prob-
ably extinct. Last specimen taken February 1892 on the Kohala
Mountains.

Genus Pseudonestor

Pseudonestor Rothschild, Bulletin British Ornith. Club, 1, 1893, p. xxxv.
Type, by monotypy, Pseudonestor xanthophrys Rothschild.

Pseudonestor xanthophrys Rothschild

Pseudonestor xanthophrys Rothschild, Bulletin British Ornith. Club, 1, 1893,

p. 36. (Maui.)
ParroUbilled Koa Finch

Range. Maui; confined to Haleakala above 5000 feet. Rare.

Genus Psittirostra

Psittirostra Temminck, Manuel d'Ornithologie, etc.', 1, 1820, p. 70. Type, by
monotypy, Loxia psittacea Gmelin.

This name has often been emended to Psittacirostra; the original
spelling is, however, as above.

Psittirostra psittacea psittacea (Gmelin)

Loxia psittacea Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 844. (in insulis
Sandwich = Hawaii.)

Psittacirostra psittacea oppidana Bangs, Proceedings Biol. Soc. Washington,
24, 1911, p. 30. (Molokai.)

Examination of large series has convinced us that characters alleged to differen-
tiate this form from psittacea (paleness, size) fall within the range of
individual variation.

Dysmorodrepanis munroi Perkins (see list of hypothetical forms).

Range. Forests of Molokai, Lanai, Maui, Hawaii below 5000 feet.
Now very rare, possibly extinct on Lanai.

BRYAN AND GREEXWAY : HAWAIIAN' BIRDS 135

?PsiTTIR0STRA PSITTACEA DEPPEI Rothschild

PsiUirostra psittacea deppei Rothschild, Bull. Brit. Ornith. Club, 15, 1905-,
p. 45. (Oahu.)

Oahu Ou

Psittacirostra psittacea (Gmelin) (part) in Wilson and Evans, Aves Hawaiiensis,
p. 80; PsiUirostra olivacea Rothschild in Rothschild, Avifauna Laysan
Island, etc., p. 193; W. A. Bryan, Key Birds Hawaiian Group, p. 54
(note); Henshaw, Birds Hawaiian Islands, p. 66; Perkins, Fauna Ha-
waiiensis (Aves), p. 435.

Range. Formerly forests of Oahu, now extinct.

There is great individual variation in adults of the populations of this
bird on every island. Particularly, there is a tendency toward albinism
noticeable in small populations. Rothschild describes this form as
differing from others in having the middle of the breast, belly, feathers
of the tibia and under tail coverts buffy whitish, in having a shorter
wing. In view of the fact of this known tendency toward albinism and
that Rothschild's own measurement for the wing of deppei (3.75
inches = 97 mm.) falls exactly within the size range of psittaeea
(95-100 mm.), we think the validity of this subspecies is questionable.

Genus Loxioides — Palila

Loxioides Oustalet, Bulletin Soc. Philom. Paris, ser. 7, 1, 1877, p. 99. Type, by
monotypy, Loxioides bailleui Oustalet.

Loxioides bailleui Oustalet

Loxioides bailleui Oustalet, Bulletin Soc. Philom. Paris, ser. 7, 1, 1877, p. 100.

(Hawaii.)
Palila
Loxioides bailleui Oustalet in all works relating to Hawaiian birds.

Range. Upper forest zones of Hawaii 4000 to 7000 feet in Kona and
Hamakua districts. Usually in or near mamane trees (Sophora).

Genus Rhodacanthis — Koa Finch

Rhodacanthis Rothschild, Annals and Magazine Nat. Hist. (6), 10, 1892, p.
110. Type, by subsequent designation, Rhodacanthis palmeri Rothschild.

136 bulletin: museum of comparative zoology

Rhodacanthis palmeri Rothschild

Rhodacanthis Palmeri Rothschild, Annals and Magazine Nat. Hist. (6), 10,

1892, p. 111. (Kona, Hawaii.)
Orange Koa Finch

Range. Above 4000 feet in mountain forests of Kona and Kau
districts, Hawaii. Usually in Koa trees (Acacia Koa). Now very rare,
possibly extinct. Last specimens collected (1892 or 1893) by Perkins.

?Rhodacanthis flaviceps Rothschild

Rhodacanthis flaviceps Rothschild, Annals and Magazine Nat. Hist. (6), 10'

1892, p. 111. (Kona, Hawaii.)
Yellow-headed Koa Finch

Range. Presumably the same as Rhodacanthis palmeri with which
it was associated when collected by Palmer for Rothschild.

These specimens differ in having yellow, not red, heads and in being
smaller (wing 94-95 mm.). No specimens have since been found.
Perkins (Fauna Hawaiiensis (Aves), pp. 436-437) suggests that these
are not two distinct species but that only one dimorphic form is in-
volved. Since the genus is now very rare it is probable that this
problem can never be solved.

Genus Telespyza

Telespyza Wilson, Ibis, 1890, p. 341. Type, by monotypy, Telespyza cantans
Wilson.

Telespyza cantans cantans Wilson

Telespyza cantans Wilson, Ibis, 1890, p. 341. (Midway Island in error =

Laysan Island.)
Lay san Finch
Telespyza flavissima Rothschild, Annals and Magazine Nat. Hist. (6), 10,

1892, p. 110. (Laysan.)

Range. Formerly on Laysan Island, where now extinct. Midway
Island, where introduced.

Telespyza cantans ultima W. A. Bryan

Telespiza ultima W. A. Bryan, Auk, 34, 1917, p. 71. (Nihoa.)
Nihoa Finch

Range. Nihoa.

BRYAN AND GREEXWAY: HAWAIIAN BIRDS 137

Genus Chloridops

Chloridops Wilson, Proceedings Zool. Soc. London, 1888, p. 218. Type, by
monotypy, Chloridops kona Wilson.

Chloridops kona Wilson

Chloridops kona Wilson, Proceedings Zool. Soc. London, 1888, p. 218. (Kona,

Hawaii.)
Kona Finch, ? Palila

Range. Kona district of Hawaii, in mountain forests from 3500 to
5500 feet. Never abundant, now very rare, perhaps extinct.

Family MELIPHAGIDAE
Genus Moho

Moho Lesson, Traite d'Ornithologie, livr. 4, 1830, p. 302. Type, by monotypy,
Merops fasciculatus Latham = Moho nobilis Merrem.

Mohohina Mathews, Bulletin British Ornith. Club, 45, 1925, p. 93. Type, by
original designation, Acridocercus bishopi Rothschild. "Differs from
Moho Lesson in having plumes on the side of the face" (Mathews).

Pseudomoho Mathews, Bulletin British Ornith. Club, 45, 1925, p. 93. Type, by
original designation, Mohoa braccatus Cassin. "Differs from Moho Lesson
in having quite a different tail formation and in not having any orna-
mental plumes" (Mathews).

Mohornis Mathews, Systema Avium Australasianarum, pt. 2, 1930, p. 800.
Type, by original designation, Moho apicalis Gould.

Moho nobilis braccatus (Cassin)

Mohoa braccata Cassin, Proceedings Acad. Nat. Sci. Philadelphia, 7, 1855,

p. 440. (Kauai.)
O-O, A-A
Acridocercus braccatus Cassin in Wilson and Evans, Aves Hawaiiensis, p. 99;

Perkins, Fauna Hawaiiensis (Aves), p. 445.

Range. Forests of Kauai. Now rare.

Moho nobilis apicalis Gould

Moho apicalis Gould, Proceedings Zool. Soc. London, 1860, p. 380. (Owhyie

in error = Oahu.)
Oahu 0-0
Acridocercus apicalis Gould in Wilson and Evans, Aves Hawaiiensis, p. 103;

Perkins, Fauna Hawaiiensis (Aves), p. 445.

138 bulletin: museum of comparative zoology

Range. Formerly forests of Oahu, now extinct. Last specimen
collected by Herr Deppe in 1837.

Moho nobilis bishopi (Rothschild)

Acrulocercus bishopi Rothschild, Bulletin British Ornith. Club, 1, 1893, p. xli-

(Molokai.)
Bishop's 0-0
Acrulocercus bishopi Rothschild in Wilson and Evans, Aves Hawaiiensis, p. Ill ;

Perkins, Fauna Hawaiiensis (Aves), p. 445.

Range. Forests of Molokai. Now rare.

Moho nobilis nobilis (Merrem)

Gracula nobilis Merrem, Avium rar . . . Icones . . ., 1, 1786, p. 7. (Hawaii.)

(fide Davies-Sherborne)
0-0 '
Acrulocercus nobilis Merrem in Wilson and Evans, Aves Hawaiiensis, p. 105;

Perkins, Fauna Hawaiiensis (Aves), p. 445.

Range. Formerly forests of Hawaii. Now probably extinct.

Genus Chaetoptila

Chaetoptila Sclater, Ibis, 1871, p. 358. Type, by original designation, Entorniza
? angustipluma Peale.

Chaetoptila angustipluma (Peale)

Entorniza angustipluma Peale, U. S. Exploring Expedition, 8, 1848, p. 147.

(Hawaii.)
Kioea (?)

Range. Formerly mountain forests of Hawaii. Now extinct.
Recorded from "between the lower Volcano House and the crater of
Kilauea" where Mills collected his specimens (fide Rothschild who
quotes Palmer in Birds Laysan Island, etc., p. 216).

"District of Hilo near Olaa" (fide Wilson and Evans, Aves Hawaii-
ensis, p. 114).

. . . "supposed to have come from Olaa, it is very doubtful whether
the Chaetoptila was found there. It is much more probable that it
was confined to the high plateau between the mountains and the
upper edges of the forest bordering this, where it was observed by
Pickering and Peale of the U. S. Exploring Expedition" (Perkins,
Fauna Hawaiiensis (Aves), p. 445). All these speculations are open
to question.

BRYAN AND GREENWAY: HAWAIIAN' BIRDS 139

Part II. HYPOTHETICAL FORMS AND RECORDS

Family ARDEIDAE— Herons

Demigretta sacra (Gmelin)

Ardea sacra Gmelin, Systema Naturae, 1, pt. 2, 1789, p. 640.

Range. Asiatic coasts and islands of the Pacific.
Recorded in the Hawaiian Group from Kahalui, Maui (a single sight
record by Finsch).

Family LARIDAE— Gulls

What is now thought to be a hybrid Larus hyperboreus Gunnerus x
Larus argentatus vegae Palmen is recorded as Larus nelsoni Henshaw
(Auk, 1, 1SS4, p. 250) by Oberholser (Auk, 35, 1918, p. 349) from
Hilo, Hawaii.

The specimen is said to be a young female and is in the U. S. National
Museum, Washington.

Subfamily DREPANIDINAE

Paroreomyza perkinsi (Rothschild)

Oreomyza perkinsi Rothschild, Avifauna Laysan Island, etc., pt. 3, 1900, p.

129. (Puulehua, Hawaii.)
Perkins's Creeper

Range. Known from a single specimen, the type, now in the
American Museum of Natural History, New York.

This specimen is intermediate in color and form of bill and is, in
our opinion, a hybrid Chlorodrepanis virens x Paroreomyza mana.
Although the feeding habits of these two birds differ, we do not think
that this would prevent occasional breeding, Perkins to the contrary
(Fauna Hawaiiensis (Aves), p. 417).

Genus "Sassius" Rothschild and Hartert

Sassius Rothschild and Hartert, Bulletin British Ornith. Club, 46, 1926, p. 51.
Type, by monotypy, Sassius simplex Rothschild and Hartert.

"Sassius simplex" Rothschild and Hartert

Sassius simplex Rothschild and Hartert, Bulletin British Ornith. Club, 46
1926, p. 51. (Sandwich Islands.)

140 bulletin: museum of comparative zoology

This curious little skin was discovered by Dr. Hartert in the
Naturhistorisches Museum in Vienna. He thought it to be an un-
described Drepanid at the time, but since then other ornithologists
have thought it to be more probably an artefact made of the skins
of Sun Birds (Nectariniidae), an African and Asiatic family (see
Meise, Proceedings 8th Internat. Orn. Congress, 1934, p. 123).

Genus "Dysmorodrepanis"

Dysmorodrepanis Perkins, Annals and Magazine Nat. Hist. (9), 3, 1919, p. 250.
Type, by monotypy, Dysmorodrepanis munroi Perkins.

"Dysmorodrepanis munroi" Perkins

Dysmorodrepanis munroi Perkins, Annals and Magazine Nat. Hist. (9), 3,
1919, p. 251. (Kaiholena Valley, Lanai.)

This curious bird is, in our opinion, an aberrant specimen of
Psittirostra psittacca (see Greenway, Auk, 56, 1939, p. 479). The type
and only specimen is in the B. P. Bishop Museum, Honolulu.

THE GENERIC ARRANGEMENT OF THE DREPANIDINAE

By James C. Greenway, Jr.

The inter-relationships of the genera of Drepanidinae have by
some been considered solved by the work of R. C. L. Perkins Fauna
Hawaiiensis (Aves) whose arrangement Sharpe followed In his Hand-
List. Perkins' arrangement, however, is not entirely satisfactory,
for in some cases he has used characters which form what appear to
be, even on the evidence he himself uses, rather unnatural groups.

There are no morphological characters to prove that these genera
are properly placed in a single subfamily, and Perkins' and Gadow's
works, which are classical, are at the same time not convincing, since
these birds have no single character in common. However, the ex-
tremely divergent genera are connected by intermediates, and the
hypothesis cannot be disproved. Whether the group sprang from a
"Coerebine" or a "Tanagrine" stock, or both, is not known. It is
therefore useless to speculate about the specialization of forms.

Within the subfamily we have two groups which may be called
"Coerebine" and "Tanagrine", simply because they resemble super-
ficially these groups. The first have tubular tongues with well devel-

BRYAN AND GREENWAY: HAWAIIAN BIRDS 141

oped brush-like tips. This group includes Viridonia, Palmeria,
Himatione, Vestiaria, Drepanis, Hemignathus, Heterorhynchus,
Loxops and Chlorodrepanis. At this point there is a sharp demar-
cation line, for the tongues of the following genera are not tubular,
those of Paroreomyza and Ciridops being concave and with a modified
brush (from this point of view intermediate). Through Psittirostra
there is gradation to the fleshy tongues of Loxioides, Rhodacanthis
and Telespiza, which are the markedly "Tanagrine" types. In spite
of the sharp difference in the shape of the tongue there is no other
character which indicates a break in the continuity of intermediate
forms, Chlorodrepanis and Paroreomyza being so close that, were it
not for the tongue, they would be considered to be congeneric under
present concepts of generic limits. Loxops, too, is intermediate,
having the bill and general form reminiscent of a goldfinch but with
a typically Drepanine tongue From two points of view this, is rather
an aberrant genus, for the shape of bill and tongue do not conform
as in other genera, and the bill with its "loxian twist" is of course
very curious. According to Perkins, it is a convergent parallelism of
both habit and form.

Perkins' arrangement groups Drepanis, Vestiaria, Ciridops, Palmeria
and Himatione together, and this assemblage he characterizes as
follows: "The genera of the first group are characterized by the
truncate apices of the primaries, except in the anomalous Palmeria,
and by the plumage of the young which is always partly black or of
a dull colour." This is an unnatural grouping and cannot be accepted.
A careful examination of these forms reveals the fact that only
Vestiaria and Himatione have truncated primaries. Furthermore,
adults of Telespiza ultima are melanistic and the melanism in cantons
is quite as impressive as in Vestiaria, Perkins to the contrary. Melan-
ism, in any event, is a character of doubtful value, such widely diver-
gent groups as Coereba and Charmosyna being classic examples of
melanistic mutation.

Although no linear arrangement is satisfying, it would appear to
be much more natural to place the tubular ("Coerebine") and the
more fleshy tongued ("Tanagrine") Drepanidinae at opposite ends of
the list, with the intermediate genera, Chlorodrepanis, Loxia, Parore-
omyza and Ciridops between them. Hemignathus is distinctly
"Coerebine" and should be placed close to Drepanis. Ciridops is of
course a very curious bird. The tongue is, however, intermediate,
and even though the color pattern (black wings and tail) resembles
Vestiaria slightly, as Perkins remarks, it would probably be better

142 bulletin: museum of comparative zoology

placed as an intermediate rather than with the typical Drepanidinae.
Ideally, Chlorodrepanis might head the list, leading us from the
Coerebinae into the Drepanidinae.

As in the "Coerebine" forms, so in the "Tanagrine" forms we are
gradually led from a tongue which in Pseudonestor approaches a
tubular form through Psittirostra, in which the Drepanine form is
still observable, to Loxioides, Rhodacanthis, Telespyza and Chloridops.
I have not been able to find any information about the tongues of the
last two genera, but Gadow places them (p. 246) near Rhodacanthis
which, he remarks, has the "most compact" tongue of all. There is
no doubt about the fact (which Gadow points out) that the narinal
structure follows, in effect, a parallel course.

It may well be argued that the last four genera should be lumped
in a single genus, as Rothschild suggests. Loxioides is the oldest name.
Lack of information about the tongues of these genera deters me
from this course.

No new facts are presented here, the anatomical information having
been presented by Gadow (in Wilson and Evans, Aves Hawaiiensis),
Rothschild and Perkins. I have examined tongues of Paroreomyza,
Chlorodrepanis, Himatione and Psittirostra. The interpretation of
evidence collected by Perkins, since accepted, seems to be faulty,
and a, to me, more natural linear arrangement, suggested by Gadow,
follows :

Viridonia

Palmeria

Himatione

Vestiaria

Drepanis

Hemignathus

Chlorodrepanis

Loxops

Paroreomyza

Ciridops

Pseudonestor

Psittirostra

Loxioides

Rhodacanthis

Telespyza

Chloridops

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 3

OBSERVATIONS ON CHINESE GOMPHINE
DRAGONELIES

By James G. Needham

CAMBRIDGE, MASS., U.S.A.

PRINTED FOR THE MUSEUM

June, 1944

PUBLICATIONS
OF THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
investigations by the Staff of the Museum or of reports by spec-
ialists upon the Museum collections or explorations.

Of the Bulletin, Vols. I to XCIII, and Vol. CXIV, No. 1, 2
have appeared and of the Memoirs, Vol. I to LVI.

After 1941 no more Memoirs are to be published.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 3

OBSERVATIONS ON CHINESE GOMPHINE
DRAGONFLIES

By James G. Needham

CAMBRIDGE, MASS., U.S.A.

PRINTED FOR THE MUSEUM

June, 1944

No. 4 — Observations on Chinese Gomphine Dragonflies
By James G. Needham

Chinese dragonflies first caught my attention in 1927, when I
received an unanticipated invitation to spend a year in China visiting
and conferring with departments of Biology in the Universities of that
country under the auspicies of the China Foundation for the Promo-
tion of Education and Culture. I looked about for aids to the study of
the local dragonflies and found there were none. There were only bare
descriptions of the adults of many species, printed in half a dozen
languages, and well scattered through the zoological literature of the
world. Nothing was known of the immature stages.

Since I was invited to lend what aid I might to the study of biology
in China, I conceived the idea that by supplying a manual for the
study of the one group of insects with which I already had some prac-
tical acquaintance at home, I might help Chinese students in the study
of their local fauna. Indeed, I was quite sure that such aid might be
more real and the results more lasting, than any that might come from
merely lecturing to them. So I began at once gathering together the
materials for a manual, going in the field to collect dragonflies as I had
opportunity, and enlisting the aid of local collectors wherever possible.

I started with no collection at all, and with almost no personal ac-
quaintance with the Oriental Odonate fauna. The literature of the
group was largely lacking in that country. Wherefore, such time as
could be taken from teaching and conferences, I devoted to collecting
and studying dragonflies.

After my arrival in September I did a little collecting of autumnal
species in and about Peiping, with the invaluable aid of Dr. Chen-fu
Wu of Yen Ching University, and Dr. Chung-lo Liu of Tsing Hua
University, and their advanced students. They guided me to the best
collecting places in some of the most beautiful aquatic situations in
North China. During the winter I was able to work on my collections
in the private laboratory of my good friend, Dr. N. Gist Gee, who was
himself a lifelong student of the Chinese fauna, and a distinguished
specialist in fresh-water sponges. In the spring he went with me to the
Yang-tze valley; to Soochow and to Hang Chow Universities where
other generous collaborators were found. I spent the month of April
in Nanking; that was my real harvest season. In Nanking I had the
generous assistance of Dr. C. Ping and a number of his research stud-
ents. There I had considerable time for collecting adult dragonflies,
and for working out partial life histories. In May I returned to Peip-

146 bulletin: museum of comparative zoology

ing, and thereafter^war conditions prevented further field work. It
was with profound regret that I had to leave China at the very opening
of the best collecting season. Then I returned home to America, bring-
ing all the collections, made and borrowed. Acknowledgment is made
in the Manual of the many sources of the borrowed material.

The Manual was written at my home in Ithaca, in such intervals
of time as I could take from teaching and departmental administra-
tion. I wrote it for the use of Chinese students in the study of their
homeland fauna. I sought merely to provide them with concise de-
scriptions, keys and tables for families, genera and species; and in all
the larger genera, where species are difficult of identification, to pro-
vide figures of the genital characters that are the ultimate criteria for
species.

In matters of classification I considered it in the interest of students
that I should keep to the older and simpler family groupings, rather
than use the many recently introduced and still untested subdivisions
of the families, concerning the validity of which the specialists are not
as yet in entire agreement. I wanted to provide something that the
college students could use; and I have had the satisfaction of knowing
that they have used the Manual successfully.

In only one group, the Gomphinae, did I add any considerable num-
ber of new species. Seventeen of these were in the large and hetero-
geneous genus Gomphus of the older authors. Among them were new
heterogeneities that I could not fit to some of the subdivisions of that
genus that had been recently proposed. I contemplated further work
upon them when more adequate collections and more time should
make that possible; but little new material has come to hand. On
completion of the Manual borrowed specimens were returned to their
owners ; but I kept duplicates of the few species that were represented
by more than one specimen; and I took occasion, while the others were
in my hands, to make photographs of the wing venation of most of
them. And now that, in retirement, I have time for more adequate
study, I have only this scanty material available. A careful restudy
of the wing venation of the Chinese Gomphines is hereinafter at-
tempted.

The venation of the basal half of the hind wing appears not to have
been thoroughly explored; for in it there are structural characters
whose taxonomic value has been quite overlooked. The late lamented
E. B. Williamson discovered some new characters in these parts, and
used them to good purpose in his well known Burmese paper of 1907.
There are yet other unused characters to which I want to direct at-

NEEDIIAM: CHINESE GOMPHINE DRAGONFLIES

147

tention here. To them I will apply convenient terms for use in the
descriptions that are to follow. The new terminology will be merely
supplemental to that used in my Manual and fully illustrated in
Plate I of that volume.

In the accompanying figure is shown a diagram based on a careful
drawing of the base of the hind wing of Gomphus campestris Ndm.,
labelled to supply names for the parts hereinafter used. The principal
longitudinal veins, Costa, Subcosta, and Radius, Media, Cubitus and

Cu, en,

Fig. 1. Base of the hind wing of Gomphus (? Burmagomphus* campestris Ndm.

Anal, are labelled in the figure at both ends of each vein, with appended
numerals designating the branches of some of them; four important
cross braces, nodus (n), subnodus (sn), arculus (ar), and anal crossing
(Ac) and three other parts, middle fork (Mf), anal loop (AL), and
membranule (to), illustrated in Plate I of the Manual, are repeated in
this diagram.

New designations here introduced are as follows: intermedian cross-
f ins (i. med.: in this wing there is but one) lie in the space between
veins M3 and M4 and beyond the arculus to the middle fork (Mf).

In the anal area of both wings the cells that lie in line alongside the
anal vein from base to gaff (g)1 are called paranals. In the fore wing
there is oftenest but a single row of them: there is always more than

1 The gaff is the fused portion of veins C'u2 and Al.

•

148 bulletin: museum of comparative zoology

one row in the hind wing and the cells of the first row are highly
differentiated and are of great systematic importance. In our diagram
they are labelled n, o, p, and AL. Cells n and o are constantly present
as large single cells in all Gomphinae known to me, with the single
exception of Anormogomphus ; cells p and AL may be divided by cross-
veins. An anal loop (AL) is said to be present only when this area is
definitely bounded by a strong vein in the rear (with that boundary
generally much stronger than shown in this figure); this loop is often
enlarged and divided into several cells.

For the cell rows running in the opposite direction, from front to
rear, (more especially for the single row alongisde vein Al extending
from the hind angle of the triangle to the wing margin and numbered
1, 2, and 3 in the diagram) I use Williamson's name post anals.

The areas into which the broad anal field of the hind wing is divided
by the three branches of the anal vein may be designated as the first
(x), second (y) and third (z) anal interspaces. Each lies behind the
branch bearing its own number (principal veins being numbered
around the wing margin from front to rear). These interspaces differ
in breadth and slant, and in number, size and arrangement of the cells
composing them. Though little noticed hitherto, they offer excellent
systematic characters. The third interspace (z) is modified into the
anal triangle of the male.

GOMPHUS s.lat.

I wish now to make a further analysis of the species that I lumped
together in the great genus Gomphus in the Manual. As evidenced by
wing venation, these species fall into natural groups as follows:

Group 1. G. abdominalis only

For it I propose the generic name GASTROGOMPHUS. Its
characters are: a very long, thick abdomen, about a third longer than
the hind wing; anal vein 3 arises generally after, and sometimes
opposite the anal crossing; no basal subcostal cross vein, and no cross
veins in any of the triangles; first and fifth antenodals thickened; a
single row of large paranal cells in the fore wing; anal triangle of the
male three celled, and four postanals cells in the hind wing (see
Manual. PI. I, fig. 4); appendages of the male of about equal length
and divergence (Manual. PI. VI, fig. 2).

NEEDHAM: CHINESE GOMPHINE DRAGONFLIES 149

It should be noted also that in the one known species there is a very
wide differentiation in size among the cells of three wing areas; very
large before the level of the areulus, a little smaller outward to a line
drawn from the stigma to the hind angle of the wing, and much smaller
thence outward to the margin.

The nymph (Manual. PI. VII, figs. 1 and la) differs from all known
related forms in having neither dorsal hooks nor lateral spines; in
having the front border of the median labial lobe doubly produced
(bilobed) and fringed at the sides of a bare median notch; and in
having the strongly incurving terminal third of the lateral lobe very
feebly denticulate on its concave inner margin.

Group 2. Xenogomphus, gen. now Type G. agricola Selys.

Characters: middle fork (Mf) unsymmetrical, askew forward; gaff
as long as or usually longer than the inner side of the triangle; inter-
median crossveins 3/1 in fore and hind wing respectively; anal triangle
of the male hind wing usually of five cells; no basal subcostal antenodal
crossvein; first and fifth antenodals thickened; no anal loop, but
usually two complete rows of cells in the first anal interspace; male
caudal appendages of equal length but the branches of the inferior
much more widely outspread (see Manual. PI. VI, figs. 6 and 6a).

Here belong also G. succumbens Ndm. (Peking Soc. N. H. Bull. 5, 3,
1930), G. citimus Ndm., G. lautus Ndm., and probably also (judging
by similarity in form of male appendages) G. svrn-hcdini Sjostedt from
Szechuan and G. ekichibui Fraser and G. mclampus Selys from Japan.
This is the only group of species in the Gomphus series of Williamson
that has the middle fork (Mf) unsymmetrical.

The nymph (see Manual PI. VII, figs. 2 and 2a) is depressed with
strictly lanceolate abdomen bearing short triangular lateral spines on
segments 7 to 9, and low dorsal hooks on segments 3 to 9, the latter
very small at the front and regularly increasing in size to rearward to
the 8th segment. The middle lobe of the labium is prominent, tri-
angularly produced with a pair of little teeth at its slightly truncate
tip, and fringes of marginal hairs at either side. The terminal fourth
of the lateral lobe beyond the base of the strong movable hook is
roundly incurved to meet the denticulate inner border, without
forming a distinct terminal hook.

The type species of the two preceding genera are both pond species,
common in central China, where I made rearings of both of them
repeatedly.

150

bulletin: museum of comparative zoology

Group 3. Eogomphus, gen. nov. Type Gomphus neglectus Ndm.

Characters : Triangles of both fore and hind wings long in the axis
of the wing, and generally four-sided by failure of the long sides to
meet at the outermost angle ; both usually traversed by a single cross
vein; bridge vein shortened distally, the distance from subnodus to
oblique vein being about a third of the distance of subnodus from the
middle fork; gaff nearly as long as the inner side of the hind wing
triangle; basal subcostal crossvein present in fore wing, absent in the
hind; vein A2 weak and angulated so as to be almost unrecognizable;

Fig. 2. The wings of male (basal part) and female Eogomphus neglectus Ndm.

for a considerable distance from the wing margin the paired long veins
inclose more than one cell row, the greatest doubling between M3 and
M4, less between Oil and Cu2, least between Rs and M2; behind
Cu2 in the fore wing are two or three cell rows traversed by ill de-
veloped accessory branch-like sectors; and middle fork (Mf) sym-
metrical.

This genus is perhaps nearest to Davidius of the Gomphus series, but
it differs in having the fore wing triangle longer and not angulated on
the outer side; the intermedian crossveins are little reduced, being 5/2
in fore and hind wings respectively. This character transgresses the
lines heretofore drawn between the Gomphine and Epigomphine
series, as does also the general aspect of the rather elongate wings.

The nymph is unknown. The nymph figured in the Manual on
Plate VII, figures 3 and 3a, referred to on page 67 as possibly belonging
to this species, is the nymph of Merogomphus.

NEEDHAM: CHINESE GOMPHINE DRAGONFLEIS 151

The three teneral specimens representing this most unusual Gom-
phine came to me just before the manuscript of the Manual left my
hands. One male was retained for the Cornell University Collection.

After a reexamination of it, I may here add another note of de-
scription.

The face and top of the head and dorsum of the first abdominal
segment are densely hairy. The dorsum of the bilobed occiput is
thinly clad with short hair that is parted in the middle and outspread
flat both ways therefrom. The spines on femora and tibiae are very
numerous, short and in uneven alignment in the outer row on the hind
femur, with the last one in the row but little longer than the others;
those of the middle femur are more than twice as long, equally num-
erous and they form an even regular row.

The superior caudal appendages are widely divergent from the base,
tapering and convergent only toward their tips. Each ends in a small
black tooth. Below the base is a large inferior branch that ends in a
blunt black tooth. The inferior appendage is little more than half as
long as the superiors and its tips have less than half their spread. It
is quadrangular, with a straight hind margin from the angles of which
arise two stout branches that project straight to rearward. Each ends
in a blunt black upturned tooth. The genitalia of the second segment
are rather prominent. The anterior hamule somewhat resembles the
cheliped of a craw-fish with both tips bluntly rounded, the anterior tip
slightly longer and inflexed around the other. The posterior hamule is
perhaps twice as long as the anterior. There is a bulbous enlargement
of its upper third, bearing on its inner side a cluster of about a dozen
small black denticles; then suddenly tapering to a claw-like incurved
tip. The peduncle ("vesicle") of the penis slopes down to rearward,
and is deeply cleft on the anterior side for the reception of the greatly
expanded penis tip. It is nearly bare except for the edges of this cleft
and the hood-like inner side of it. The second joint is clavate toward
its tip, and lacks an apical spine. The reflexed third joint bears a re-
markable enlargement at its tip; a deeply cupped expansion that
carries a suggestion of likeness to an irregular flower. Above, the rim
of the cup is deeply emarginate, and within it arise two petal-like lobes.
Far out from its center projects a bifid stigma-like process ending in
a pair of blunt, recurved, flabellate tips, below which projects a spine
of half their length.

The apical carina of the tenth abdominal segment is produced to
rearward in a low bare triangular prominence, on either side of which
is the usual line of black denticles.

152 bulletin: museum of comparative zoology

Group 4. Merogomphus Martin. Type M. paviei Martin

A more careful examination of the single known female specimen of
Gomphus torpens Ndm. of the Manual leads me to conclude that it
should have been placed in this genus and associated with Merogomphus
vandykei Ndm., for it has the following characters : vein A3 arises op-
posite the anal crossing (Ac), and not beyond it; basal subcostal cross
vein (an) present in the fore wing; intermedial! crossveins 3/1 in fore
and hind wing respectively, middle fork (Mf) symmetrical; gaff more
than half as long as the inner side of the triangle ; two rows of paranal
cells in the fore wing; first anal interspace (x) wider than the second
(y), and no anal loop. It also has a half a dozen very large spines in the
outer row on the hind femur.

A nymph of this genus was figured in the Manual without name
(Plate VII, figs. 3 and 3a) and described on pages 66 and 67. It is
probably the nymph of M. vandykei, as determined by a recent study
of the venation of its crumpled wings.1

Group 5. Mesogomphus Foerster. Type Gomphus cognatus Rambur

The Gomphus brevipennis Ndm. of the Manual belongs here, as
evidenced by the following characters: vein. A3 arises just before or
opposite the anal crossing (.4c); two rows of paranal cells in the fore,
wing; intermedian crossveins 2/1 in fore and hind wing respectively;
first and fifth antenodals thickened; middle fork (Mf) symmetrical;
arculus unusually close to the triangle of both fore and hind wings and
the front side of the subtriangle much shorter than the inner side of
the triangle; four postanal cells in the hind wing; four or five cells in
the male anal triangle; no anal loop, and the first anal interspace
wider than the second.

The nymph has been described and figured for one species of this
genus, M. balneorum by Needham and Gyger (Philippine Jour. Sci.
63, 33,P1.X, figs. 125 and 126) and for two others, M. lineatus and M.
reinwardti by Lieftinck (Tijd.v.Entomol. 77,21, 1934).

Group 6. Burmagomphus Williamson. Type B. williamsoni

Fraser

The Gomphus dolus Ndm. of the Manual belongs here. In venation
it is very close indeed to the type species as figured by Williamson

1 The method used was that described by Dr. May K. Gyger in Entomological News, 50,
p. 21, 1939.

NEEDHAM: CHINESE GOMPHINE DRAGONFLIES 153

(Proc. U. S. Nat. Mux. 33, 298, fig. 27, 1907). The combination of
venational characters by which this genus has been set apart is as
follows: In the fore wing, a single row of paranal cells; a small triangle
slightly angulated near the middle of its outer side; a long close
parallel of veins M4 and (Ail beyond it with but two intervening
rows of cells out to the level of the oblique vein, with a rather sudden
widening thereafter; and in the hind wing but three postanal cells.

The nymph of this genus has been mentioned, and given an un-
intelligible two-line description by Fraser in Fauna of British India:
Odonata 2, 212, 1934.

Another small species that would appear to belong near to Burma-
gomphus is the one a portion of whose hind wing is shown in the
accompanying figure of Gomphus campestris Ndm. of the Manual.
(See figure 1.) It is of small size (hind wing 25 mm), with slightly
angulated outer side to the hind wing triangle, and only three postanal
cells. The single-celled anal loop is quite like that of B. icilliamsoni.
There are these discordant characteristics: there is a basal subcostal
crossvein in both fore and hind wings; one or two cells of the paranal
row in the fore wing are divided, and the double row of cells beyond the
triangle does not extend outward beyond the level of the nodus.1
The venation as a whole is more sparse, there being but 11:7/8:7
nodal crossveins in fore and hind wing respectively and only three
crossveins under the bridge.

My material is inadequate for determining whether these differ-
ences are constant enough to justify generic separation. Since this
species has been taken on the campus of Yen Ching University, it
should be possible for some one there to obtain additional specimens
including also its immature stages.

Group 7. Gomphus s. str. Type Libellula vulgatissima Linn

The remaining species appearing under this generic name in the
Manual may be allowed to remain so for the present. They show a
general conformity to the type, but with numerous small divergencies
which I shall now try to indicate in so far as they appear in the vena-
tion of the wings. They all seem to have the following characters in
common: middle fork (Mf) symmetrical; vein A3 arises beyond the
anal crossing (Ac); intermedian crossveins generally 2/1; paranal
cells in the fore wing more than a single row, some cells at least being

1 This disagreement applies also to Fraser's figure of Burmagomphus pvramidalis, Faun.
British India Odonata: 2, 212, fig. 66, 1934.

154 bulletin: museum of comparative zoology

divided; postanal cells four to seven; anal triangle of the male gener-
ally three celled; and no basal subcostal cross veins.

Recalling Burniagomphus are the two small shortwinged species
G. arvalis Ndm. and G. sowerbyi Ndm. These have a well defined one
celled anal loop with the base of vein Al kinked around its outer
corner. The outer side of the nearly equilateral fore wing triangle is
slightly angulated in the middle. The arculus is between the first
and second antenodal cross veins. The gaff is about half as long as
the inner side of the hind wing triangle. Vein M4 is slightly undulate,
and there are four post anal cells. This latter character prevents plac-
ing them in Platygomphus, as does also the well developed 3-celled
anal triangle in the male. De Selys placed a question mark before his
Platygomphus occultus when he placed it in that genus. It belongs
rather with the above named pair. The three might possibly be made
the basis of a new genus; but until Burmagomphus and Platygomphus
are better defined, and until Gomphus campcstris has found its place,
and until the nymphs of all of them are made known, another name
would only add to the confusion.

Two somewhat larger species of the Manual, G. intinctus and G.
collar is, are like the three preceding in most characters, but the
arculus is nearer the second antenodal crossvein, and intinctus has
five postanal cells. In all five the gaff is about half as long as the inner
side of the triangle.

A peculiar species that is known unfortunately from only a single
female specimen is G. edax Ndm. The triangles are both elongated in
axis of the wings, the outer end of the hind wing triangle being turned
up at the end like a sled runner (see the next figure) ; there are seven
postanal cells ; the branches of the anal vein are aslant outward.

Another peculiar species, described later by me (lAngnan Sci. Jour.
10, 227, 1931) from a single female specimen taken in Hainan is
G. hoffmani. It has the first and sixth antenodals thickened, the
arculus at or beyond the second; no basal subcostal antenodal cross-
vein, the fore wing with two rows of paranal cells, no large cells in the
anal area behind the first paranal row, and the first anal interspace
much wider than the second. Added to this array of differences there
is a peculiarly elongated three-celled anal loop, around the outer end
of which vein Al makes a short sharp bend. Also the hind wing is
widest at the nodus. The gaff is as long as the inner side of the tri-
angle, which latter in the fore wing is a little longer than the front side.
Whether this wing is quite normal I cannot say.

There remain six large species (hind wing 37-40 mm.) that conform

NEEDHAM: CHINESE GOMPHINE DRAGONFLIES 155

more closely (still none too well) to the type of the genus. One,
G. cuneatus Xdm. of the Manual appears to be distinguished by having
the veins M3 and M4 not at all undulate, strictly parallel and regularly
curved and inclosing somewhat larger cells than in the other five
species. In general it has a more open venation. It also has a longer
gaff, almost as long as the inner side of the triangle, and a shorter
kink in the base of the anal vein at the outer angle of the one celled
anal loop. There is in the type specimen but one extra cell in the
otherwise single paranal row of the fore wings.

Of the five remaining species, G. endicotti appears to be separable by
reason of the shortness of the front side of the fore wing triangle — not
longer than the inner side of the same; and G. flavicomis (Peking Soc.
Nat. Hist. Bull. 1, 2, 1930) by having its anal crossing close to the
inner end of the subtriangle in the hind wing — less than half its own
length therefrom.

Finally G. amicus is separable from the remaining two by the
shortness of its gaff — less than half the length of the inner side of the
front wing triangle; and these last two may be separated by the posi-
tion of the arculus in relation to antenodal crossveins: it is midway
between the first and second in G. clathratus, and at or very close to
the second in G. septimus.

THE FRAMEWORK OF THE WING ABOUT THE

TRIANGLES

In the preceding pages I have been pointing out the best single vena-
tional characters that I have been able to find for distinguishing each
of the species listed in my Manual under GOMPHLS (all of them ex-
cept G. sornnolens, of which I now have neither wings nor photographs
of venation available). I now wish to present in the form of a table
some correlations of characters especially to show the relations of the
parts of the strong framework of the wing that are in or around the
triangles.

As a standard of comparison I take the part marked a, which forms
the one common side of triangle and subtriangle, and which is formed
in development about the posteriorly deflected portion of the main
Cubital trachea. Two additional reasons for its selection are (1) its
central location, its ends being in contact with all the other parts
compared; and (2) its relative constancy in length. A little comparison

156

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showed all the other parts to be more variable. Next in constancy was
the part marked e, which is formed about the main Anal trachea.

The part a was given an assigned value of 10, and the length of all
the other parts were estimated in tenths of it. That is the meaning of
the numerals in the central columns of the table. These values are

Fig. 3. The parts about the triangle (T) in the wing hind of Gomphus edax Ndm. to illustrate
the terms used in the following table.

merely estimates made under inspection with a lens, without careful
measurements: wherefore allowance (possibly up to 10%) may have
to be made for errors of judgment, and additional allowance for varia-
tion in individual specimens.

The accompanying enlarged diagram, based on a drawing of the
wing of Gomphus edax Ndm. will be useful for comparison.

In general it may be said concerning venational characters that the
conjunctions and proportions and directions of the component parts
of the strong framework of the wing, and the layout of the spaces be-
tween principal veins and their branches offer far more dependable
taxonomic characters than are to be found in the number of interven-
ing crossveins.

NEEDHAM: CHINESE GOMPHIXE DRAGOXFLU SS

157

Venational Characters in 20 Species of Gomphus

Species

Hind
wing

I n ter-
med. CVS.

f/h tv.

Length of

parts about

hind W. triangle

b e d e f

extra
paran.

lis

post
a rials A3sAc

A. loop
present

abdominalis

2/1

17 18 9

11 5

opp

yes

agricola

3/1

16 18 10

11 6

out

amicus

2/1

16 18 11

12 8

yes

arvalis

2/1

18 20 12

12 7

yes

campestris

clathratus

collaris

26
38

2/1
3/1
2/1

10
10
10

12 13 11
17 19 10
15 16 10

12 5
11 7

11 7

5
5

1
2

3 •

4
4

n
n

yes
yes
yes

cuneatus

2/1

15 16 10

11 7

yes

dolus

2/1

9 11 9

11 4

edax
endicotti

35
34

3/1

2/1

10
10

18 20 11
17 19 10

12 6
11 7

8
9

2
3

5 i
5 i

yes
yes

flavicornis

2/1

17 19 10

11 5

5 ]

yes

gideon
hoffmanni

36
34

4/2
2/1

10
10

16 17 10
15 16 10

11 6

11 7

5
9

3
4

4
5

n
n

yes
no

intinctus

2/1

14 15 10

11 7

yes

neglectus

5/2

19 20 10

11 5

out

occultus

2/1

14 15 9

10 5

yes

septimus
sowerbyi

• 40
29

2/1
2/1

10
10

13 15 9

14 15 9

11 6
11 6

7
4

3
2

in
in

yes
yes

torpens

3/1

15 16 9

11 4

4 i

Column 1. Length of hind wing in millimeters.

Column 2. Number of crossveins joining the sectors of the arculus between the arculus
and the middle fork in fore and hind wing.

Column 3. Relative lengths of the parts about the triangle of the hind wing in terms of
tenths of the length of the inner side of the triangle.

Column 4. Number of extra paranal cells in the front wing (more than the single row always
present).

Column 5. Number of cells in the postanal row on the proximal side of vein 1st A between
the triangle and the hind margin of the wing.

Column 6. Position of origin of vein 3d A: in, proximal to it; opp, opposite it; out, distal
to it.

Column 7. Anal loop of hind wing.

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Leptogomphus unicornis Ndm.

A study of the wings of the single known specimen of this species
shows it to have been misplaced in the Manual in the genus Davidius.
It has more in common with Leptogomphus, including (1) the form of
the wings; (2) the lack of brace vein to the stigma; (3) the trigonal
interspace regularly widening outward to the wing margin; (4) the
small triangles ; (5) the narrow fore wing subtriangle ; (6) the little ex-
panded anal area of the wing; and (7) the 3rd anal interspace (z)
longer in the axis of the wing than wide. All these I regard as primitive
characters, of relative fixity.

Fig. 4. Wings of ? Leptogomphus unicornis Ndm.

It seems to differ from Leptogomphus as represented by its type
species, L. sauteri Selys, in having but a single row of very large
paranal cells in the fore wing, in having no basal subcostal antenodal
crossvein, and in having crossveins in all the triangles and in the
supratriangular space of both the wings. I present a figure of the vena-
tion to call attention to these discrepancies. At first glance they seemed
to me to be so great as to call for generic separation; but on further
experience with the Epigomphus alliance, I think they are very un-
reliable variants, having only specific value.

NEEDHAM: CHINESE GOMPHINE DRAGONFLIES 159

TWO NEW SPECIES AND SOME NEW RECORDS

Since the publication of the Manual and of the Additions and Cor-
rections (in the Peking Soc. of Nat. Hist. 5, 1-10, 1930) only one small
collection of dragonflies has come to me from China. It was from Dr.
Ting-wei Lew. It contained two new species of Gomphines and es-
tablished a few new records.

Gomphurus gideox spec. nov.

Length 63 mm.; abdomen 45; hind wing 38.

This is a blackish species with spatulately dilated abdomen; face
black with an oblong stripe of yellow covering about half of the lab-
rum. A similar somewhat larger isolated stripe covers the top of the
frons. The remainder of the top and rear of the head is black.

The synthorax is black in front with a pair of isolated dorsal stripes
that are divergent downward, not reaching the divided cross stripe on
the collar. An antehumeral stripe of yellow is represented by a very
small spot high up near the crest, and a thin faint streak low down on
the side, its lower end. Behind the broad black humeral band the
sides are yellow with a black line on the third lateral suture, that con-
nects with the back subalar carina above, and below runs down behind
the coxa of the middle legs. A vestige of a middle stripe is present in
front of the spiracle. Legs black beyond the short bicolored basal
segments. Wings hyaline, with a faint tinge of yellow in the mem-
brane. Ante- and postnodal crossveins are 16:12 and 10/12 in fore
and hind wing respectively.

The abdomen is very moderately enlarged on the two basal segments
narrowly cylindric on segments 3 to 7, and spatulate on 7 to 10, with
widely flaring lateral expansion of the margins of 7, 8 and 9. The
dorsum of 1 is mainly yellowT and a narrow lanceolate spot appears on
the base of 2. The sides of 1 and 2 are mainly yellow, and also the base
of 3. Basal yellowish rings on 3 to 7 become narrower to rearward,
with only fine yellow intersegmental lines across the apices of 7, 8,
and 9: segment 10 and appendages wholly black. The relative length
of the last four segments middorsally is about as 11; 9: 10:5; and ap-
pendages, on the same scale as 7. Diffuse large yellow spots cover
about half of the sides of segments 8 and 9. Caudal appendages are as

160 bulletin: museum of comparative zoology

shown in figure. The posterior hamules of the male, completely hiding
the anterior ones, project strongly downward even beyond the level
of the "vesicle", and taper to claw-like sharp tips that are directed
forward.

Fig. 5. The abdomen of Gomphurus gideon sp.n., dorsal view.

The female is very similar in coloration, with the yellow areas a little
more extended, especially on the abdomen. The subgenital plate is
divided deeply into two blunt equilateral triangles that extend to
rearward across about a fifth of the 9th sternite. The supraanal plate
is shining black above, yellow beneath.

A single pair, type and paratype, collected in Chengtu, Szechuan
on June 29th, 1929, and sent me by Dr. Lew. They are now in the
Cornell University Collection.

Because of the striking dilatation of the 7th and 8th segments of the
abdomen, shown in the figure herewith, and general accord in caudal
appendages and in venation, I have placed this species in Gomphurus;
a genus hitherto known only from North America. Another Chinese
species, Gomphus kryenbergi Ris, compared by its describer with the
American Gomphus scudderi, doubtless belongs beside it. G. gideon has
however a somewhat more copious venation than the American
species, with four cells in the anal triangle in both right and left hind
wings of the male: intermedian cross veins 4/2 in both male and female,
and six postanal cells.

NEEDHAM: CHINESE GOMPHINE DRAGONFLIES 161

Davidius serenus spec. nov.

Female; length 41 mm.; abdomen 31; hind wing 27.

This is a small blackish species with yellow sides. The head is all
black except the outer sides of the mandibles and a broad transverse
stripe across the very low prominence of the frons, which are yellow.
The thorax is black in front except for a pair of opposed 7-marks that
just meet at the middle of the collar. The stalks of the 7s are slightly
tapered upward and blunt at their isolated upper ends. There is no
antehumeral yellow stripe at all. Behind the very broad black humeral
band the sides are mainly yellow with a narrow black stripe on the
third lateral suture that is connected forward with the humeral above
and below. The black of the ventral side extends upward at the middle
suture to cover the spiracle. The legs are black. The long slender hind
femora are sagged downward in the middle and beset underneath with
more than a score of slender black short subequal spines.

Wings hyaline. Ante- and postnodals 13:12/9:11 in fore and hind
wing respectively. There is an extra cubito-anal crossvein in the fore
wing, and there is a single row of cells behind the anal vein.

Abdomen mostly black with a diminishing amount of yellow on the
sides of segments 1 to 7; segment 1 mostly yellow dorsally and 2 with
a lanceolate streak of the same color. On the sides of segments 3 to 5
the yellow is broken into a row of three spots; reduced to two spots on
6, and to a single spot on 7: 8 to 10 black. Appendages yellow.

The subgenital plate of the female is oblong flat, slightly tapering to
rearward, with a deep notch at the tip, and about four fifths as long
as the venter of the 9th segment.

This species is nearly allied to D. trox Ndm., but differs in being
smaller in size, in having the labium all black, in lacking the pale spot
that is a vestige of the antehumeral stripe, in lacking the J-spot at the
rear of the side of the thorax, and in having the abdominal segments
wholly black and the appendages yellow.

There is a single specimen collected at Ruling, China in July 1933
by Dr. Ting-wei Lew, and now in the Cornell University collection.

Among the specimens sent me by Dr. Lew were five females of
Gomphus septimus Ndm. The male was described in the Manual, p. 61.
The female, heretofore unknown, is like the male in coloration, with
a basal yellow halfring on abdominal segment 7 more conspicuous than
in the male. The relative length of the last four abdominal segments
is as 15:12:10:7, with the appendages 8, on the same scale. The promi-
nent subgenital plate is scoop-shaped or shaped like the spout on a

162 bulletin: museum of comparative zoology

pitcher, triangular, black, more than half as long as the venter of the
9th segment, and directed conspicuously downward. Among the rather
stout spines on the hind femur are four to six stronger than the others,
but intermixed with the others, and none of them is as long as the
femur is thick.

Two of the specimens came from Foochow in May; one from Mt.
Poliang ding, near Ho-kiang and Ming-kiant Fukein; and two from
Tu-ching, Min-giang in Fukien.

Dr. Lew sent also a fine pair of Mcgalogomphus sommeri Selys from
Kuling; a species that has hitherto been without definite locality
assignment in China.

TWO CORRECTIONS FOR THE MANUAL AND
A CONFIRMATION

(1) Agricnemis amelia Ndm. (Manual, p. 256) is a synonym of
Ischnura delicata Hagen, which is in turn considered by some authori-
ties to be a synonym of Ischnura aurora Burmeister.

(2) Taolcstes ncctans Ndm. (Manual, p. 256) is correctly described
and illustrated, but the nymph associated with it belongs elsewhere.
The female type specimen was presented to me along with a cast
nymphal skin, supposedly reared, and with several nymphs that had
been collected at the same spot. I uncritically accepted them as be-
longing together. The nymphs were not well preserved; but in a
recent examination of one of the best of them I find enough venation
still remaining in its wing pads to show that the antenodal crossveins
are numerous. That is sufficient to show that they are not Taolcstes.

The nymph is structurally very similar to those now known belong-
ing to the genus Euphaca. Judging by its size, it should be Euphaca
opaca.

A word about the placement of Taolcstes. As explained at the be-
ginning of this article, I followed the older and easier system that
segregated the Lestinae from the others on one principal character:
middle fork (Mf) nearer the arculus than the nodus. But in doing so
I pointed out (p. 226) the nonconformity of Taolcstes with the true
Lestinae.

Two additional species have since been described from China by
Erich Schmidt (Koitowia 10, 178-183, 1931) as species of Rhipidolcstes:
R. bidens and R. truncatidens; they conform much more closely to
Taolcstes, not only in having the middle fork nearer the arculus than

NEEDHAM: CHINESE GOMPHINE DRAGONFLIES 163

to the nodus, but also in many other points of venation, and in the
male genitalia. The species T. nectans differs from both in having un-
marked hyaline wings, much more open venation, and a shorter and
thicker stigma.

Concerning the nymph of Megalestes, I noted on page 229 of the
Manual that Laidlaw had described a nymph that he referred by sup-
position to M. major (Ind. Mus. Record, 19, 185-187, 1920). I said I
was not convinced that Laidlaw's nymph belonged to Megalestes.
My remark was more than justified: this, notwithstanding Leiftinck's
oracular pronouncement (Treubia 17, 58-61, 1939) in support of Laid-
law's supposition. I obtained an almost identical, certainly congeneric
nymph of Rhinagrion philippinum from Luzon, well preserved, and
showing so complete venation in its wing pads as to leave no doubt as
to its identity. It is described and figured in the Philippine Journal of
Science, 70, 266, Plate 15, figs. 206-213 and 215-216, 1939. Laidlaw's
nymph is Rhinagrion.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 4

P3AMMOCHARIDAE

(Spider-Wasps)

Notes and Descriptions

By Nathan Banks

CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM

June, 1944

PUBLICATIONS
OF THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
investigations by the Staff of the Museum or of reports by spec-
ialists upon the Museum collections or explorations.

Of the Bulletin, Vols. I to XCIII, and Vol. CXIV, No. 1, 2
and 3 have appeared and of the Memoirs, Vol. I to LVI.

After 1941 no more Memoirs are to be published.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 4

PSAMMOCHARIDAE

(Spider-Wasps)

Notes and Descriptions

By Nathan Banks

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

June, 1944

No. 4 — Psammocharidae1

(Spider- Wasps) : Notes and Descriptions

By Nathan Banks

In continuation of previous studies on the fine collection of Psam-
mocharidae in the Museum of Comparative Zoology I present descrip-
tions of some new genera and species and notes on others. Neotypes
are made for two of Say's species and descriptions of varietal forms.
Some of these varietal forms illustrate the faunal divergence of the
New England area, which I have previously noted in a Myrmeleonid,
Hesperolcon abdominalis Say. Descriptions are given of the males of
three West Indian species previously known only from the females.
Keys are made for the species of Dipogon and for the males of the
species of Ageniella known to me from the Eastern States. A compari-
son of our large black Arizona Pepsis with a series of both sexes of the
related Cuban form and the South American Pepsis grossa shows that
our form is a distinct species.

a. NEARCTIC
Batazoxus interruptus Say

Say's diagnosis says "metathorax at tip bifasciate with yellow", and
the pleura with two yellow spots. It came from Indiana. This is the
form common in the middle states, west to Kansas and Nebraska, and
in Texas, North Carolina, etc. I have selected as neotype a female
from Chicago, collected by Prof. C. T. Brues, which agrees with the
description.

There is a form in the northeastern states which I call

Batazoxus ixterruptus var. cressoni var. now

The thorax and propodeum are black, apical margin of propodeum
with a broadly interrupted yellow line, no broad yellow band above it,
no marks on pleura, the hind margin of pronotum narrowly yellow,
and the scutellum yellow. The yellow on each side of face in the typical
form is here reduced to a slender orbital line, the clypeus is usually
black. The abdomen shows no pale at bases of segments, and the base
of first segment is usually black. The wings are darker. In the male
the yellow is likewise greatly reduced. The size is generally smaller.

The holotype is a female from Holliston, Mass., 4 August. Para-
types from Holliston from 24 June to 7 September; Wood's Hole,

'Published with the aid of a special gift from Mr. R. G. Agassiz.

168 bulletin: museum of comparative zoology

Mass., (C. T. Brues); Orient, N. Y., 12 July; White Plains, N. Y.,
Sept.; Wellesley, Mass., 13 July (Bolster); Trenton, N. J.; Penna.
(Melsheimer) ; Charter Oak, Penna, 11 August, (Kirk); and Glen-
carlyn and Falls Church in northern Virginia from 14 June to 30 July.

Type M. C. Z. No. 25729.

Pompilus ichneumonides D. T. (ichneumoniformis Patton) and P.
willistoni Patton belong to Batazonus. The latter I consider as a form
of interruptus, and the former as B. navus Cress.

Arachnophroctonus Ferrugineus Say

Say says "Ferrugineous; wings violet; pleura and metathorax
black." Inhabits U. S. The type was a male.

A specimen which agrees with the above and also with the fuller
description I have selected as neotype; it is from Falls Church, Va.;
23 July.

There are three varieties which can be separated on color; there is
scarcely any difference in structure, and the genital plate of the male
has a prominent median carina in all.

A. ferrugineus var. unicolor var. nov.

Some years ago Viereck labeled a specimen "unicolor"; it was from
the far west. The body is wholly ferrugineous except a small black
mark at the base of the abdomen ; nothing dark on thorax nor on pro-
podeum, nor even a trace of dark bands on abdomen; moreover the
wings are a yellowish brown, very much paler than those of the typical
form. The structure appears to be the same except that the clypeus is
not so deeply emarginate in the middle. The holotype is from Oak
Creek Canon, Arizona, 6000 ft., July (F. H. Snow) in the M. C. Z.
no. 25730. Others are from Tucson, and southern Arizona (Bequaert).

A. ferrugineus var. annexus var. nov.

From parts of Texas come specimens intermediate in some ways
between unicolor and typical ferrugineus. There is no black on thorax
nor on propodeum, but the abdomen above shows at least traces of a
dark band at end of first and second segments. The wings are just as
dark as in typical form, which separates it from unicolor.

Holotype is from Fedor, Lee Co., Texas, June 15, 1909 (Birkmann);
paratypes from Fedor in June and September, also Dallas, Texas
(Boll); Austin, Texas, 13 October (Brues); and Davis Mts., Texas, 28
June (Englehart). Type, M. C. Z. no. 25731.

Typical ferrugineus also occurs in Texas.

hanks: psammocharidae 169

A. FERRUGINEUS var. NIGRESCENS var. nOV.

In the northeastern part of the country there is a much darker form;
the wings are nearly black; the abdomen, in the female, shows the
dark bands at tips of first and second segments. The thorax and pro-
podeum are wholly black above and below, and on pleura; the femora
are at least partly black, the hind femora often wholly so; in the
female the tip of tibia and of some tarsal joints also black; the antenna
is nearly wholly black, basal joint only remaining pale; in the male the
basal segment may be more black than usual; in females the face
above the antennae is more or less darkened. The clypeus is less deeply
emarginate than in typical form; otherwise I note no structural
difference.

The holotype is a female from Arlington, Va., 1 1 August; paratypes
from Fort Lee district, X. J., August; Woods Hole, Mass., (Brues);
Holliston, Mass., 27 July (Bks.), and Coldbrook, Conn., 14 July
(Wheeler). Type, M, C. Z., no. 25732.

Anoplius imbellis spec. nov.

Black throughout, wings not darker at tip. Moderately long black
hair on head, thorax, and propodeum; some stiff black bristles above
tip of abdomen, and some slender curved ones above and below; no
hair on pleura. ( 'lypeus fully two and one-half times as broad as long,
below truncate; antennae slender, third joint fully five times as long
as broad, fourth about two-thirds as long as third. Face but little
narrower above than below; a median line from antennae to the an-
terior ocellus, ocelli subequal, laterals a little nearer to each other than
to eyes; pronotum deeply angulate behind; propodeum with no dis-
tinct median groove; abdomen moderately broad at base, slender and
pointed at tip; mid and hind tibiae with short spines above; long spur
of hind tibia nearly three-fifths of basitarsus; claws with short, erect
tooth.

In fore wings the basal vein a little before the transverse, marginal
cell more than length from the wing-tip, outer side straight; second
submarginal cell a little longer than broad, receiving the first recurrent
near tip; third submarginal triangular or nearly so, outer side convex,
receiving the second recurrent (lightly curved) near middle.

Length 9 7.5 to 8.5 mm.

What is doubtless the male is similar, but little more slender; the
antennae heavy, third joint about three times as long as broad; the
third submarginal cell triangular or even pedicellate, shorter than the

170 bulletin: museum of comparative zoology

second; the fourth and fifth ventral segments are covered with a
brush of black hair, denser and longer on sides ; the genitalia show a
median plate, narrow, almost pointed at tip, and with a median,
scarcely elevated ridge.

Length c? 7 to 8 mm.

Females from Corvallis, Oregon, 30 August, 4 September, and
Hillsboro, Oregon, 25 September.

Males from Corvallis, 6, 25, 30 August, and Cornelius, Oregon,
4 August (Scullen coll.).

Type in Oregon Agricultural College, paratypes there and in
Museum of Comparative Zoology no. 25734.

Nannopompilus texanus spec. nov.

Similar in structure to AT. subviolaceus. Abdomen only faintly
bluish; fore wings scarcely smoky, but a broad dark band at tip; hind
wings hyaline, smoky at tip. The body, antennae, and basal joints of
legs sericeous, strong on lower face, clypeus, coxae, and posterior sides
of propodeum. Antennae fully as short as in Ar. subviolaceus, head,
thorax, and propodeum base as in that species. Venation similar, the
second submarginal cell nearly as high as long; the recurrent veins end
beyond middle of cells. The legs also similar, except that the spines
of the comb are longer and thickened, and some a little curved, but
none plainly spatulate. Marginal cell a little more than its length from
tip of wing.

Length 7 mm.

One female from Richmond, Fort Bend Co., 22 June, (Bequaert)
and another from Fedor, Lee Co., (Birkmann), both Texas. Type
M. C. Z. No. 25S95. Separated from A:. subviolaceus by the much
paler wings, sericeous body, stouter spines.

Gymnochares texana spec. nov.

Black, pronotum slightly sericeous near hind border, and in front
near collar; antennae dark ferruginous; wings slightly darkened
throughout, tip a little darker, but not a band. Clypeus more than
twice as broad as long, lower edge truncate; each side above clypeus
the face is faintly sericeous, also the base of mandible; face with nearly
parallel sides, except near top; a very distinct median groove from an-
terior ocellus to antennae, latter slender, third joint longer than ver-
tex-width and one and a half times longer than fourth, latter but little

banks: psammocharidae 171

if any longer than fifth. Hind ocelli a little nearer the eyes than to each
other; vertex with a few fine, long hairs; pronotum almost angulate
behind, faintly depressed near hind border, and with a median groove
reaching forward; pro and mesonotum without hair, as also the pro-
podeum, latter with a very distinct median groove above; on the sides
of the posterior slope and extending over the side are distinct but not
strong transverse ridges, fading out toward center. Abdomen slightly
compressed near tip, scarcely a trace of hair above toward tip, below
a few near tip.

Legs slender as in other species; hind femora above near tip with
three well-separated minute hair-pits, rather fainter than in allied
species; hind tibia with only a few scattered minute spines, three
minute ones on apical half of outer side; inner spur of hind tibiae not
one-half as long as basitarsus, latter with only a trace of spines below.

Fore wings with marginal cell about its length from wing-tip, about
two and a half times as long as broad, not quite as slender as in birk-
manni; second submarginal cell fully one and a half times as long as
broad, receiving the first recurrent a little before tip (nearer middle in
birkmanni); third submarginal cell only a little longer than second,
narrowed one-third above, receiving the second recurrent a little be-
yond middle; basal vein before transverse; cubitus extending out to
margin as in other species. In hind wing the anal ends much before
the fork.

Length 8 mm.

One female from Austin, Texas, May (Melander), Type, M. C. Z.
25704.

Similar in general to birkmanni, but readily separated by the groove
on propodeum.

Nannochilus gen. now

Two of our species described under Ageniella, externa and congrua,
do not have a petiolate, but a sessile abdomen; they are therefore re-
lated to Priocnemis, but differ from that genus in lacking teeth or even
bristles on the hind tibia. The claws are cleft, the inner process being
very broad and obliquely truncate, and the pulvillus is extremely
large. The venation is similar to that of Pseudagenia, but the basal
vein is interstitial with the transverse. Besides Ageniella externa Bks.
which is the genotype, there is A. congrua Cress., and a new species
described herewith.

172 bulletin: museum of comparative zoology

Nannochilus osoria spec. nov.

Body black, abdomen sometimes more brown; antennae and legs
brown, sometimes the mid and hind femora are somewhat rufous, and
the front tibiae and tarsi are usually pale yellowish, all spurs pale, but
not snow-white. Clypeus and lower face with silvery pubescence, little
on vertex, but more prominent on coxae and on each side of propodeum
behind the spiracles.

Fore wings rather evenly dusky, tip not darker; veins brown to
black; hind wings more hyaline.

Clypeus broad, short, and truncate below; lateral ocelli a little
nearer to each other than to eyes; antennae moderately long and
slender, much as in iridipennis; pronotum arcuate behind, finely
granular, a median groove on basal part; abdomen with short pale hair
on venter; basal segment not nearly twice as long as broad behind.

Fore wings with submarginal cells much broader than the marginal
cell; second submarginal about as broad as long, receiving the first
recurrent near middle; third submarginal hardly longer below, but
larger, the posterior side almost angulate in middle, so the cell is no
broader below than above, receiving the second recurrent vein near
middle; basal vein interstitial with nervellus, its lower part not
strongly curved. In hind wings the anal vein ends before the fork of
cubitus.

Length of fore wing 5 to 6.5 mm.

Holotype male from Falls Church, Virginia, 12 July; paratypes from
the same locality from 28 June to 7 September; also from Great Falls,
Virginia, 20 June, and Chain Bridge, Virginia, 23 June. Type M. C. Z.
No. 25907.

Three females agree in general; dull black, sericeous, venation as in
male, spurs pale, clypeus truncate below; third antennal joint a little
longer than fourth; pronotum broadly arcuate behind; inner spur of
hind tibiae hardly one-half of basitarsus; basal segment of abdomen
broad and sessile.

Length of fore wing of female, 6 mm.

All from Falls Church, Virginia, 13 September.

Priocnemis occldentis spec. nov.

Closely related to the Eastern P. nothus; in color the wings are uni-
formly darker and the abdomen is entirely reddish, the last few seg-
ments not darkened as in nothus. The venation the same as that of
nothus even to the bend in the lower part of the submarginal cross-

banks: psammocharidae L73

vein; the propodeum is less evenly rounded than in nothus, seen from
the side there is almost an angle at the turn, the posterior slope steeper
than in nothus.

The second plus third antenna] joints not equal to vertex-width;
ocelli subequal, in a slightly broader triangle than nothus, the laterals
much nearer to each other than to eyes; pronotum angulate behind;
no median groove on propodeum, the hue striae as in nothus; the spines
and spurs on legs as in that species.

Length 6.5 to 8 mm.

From Corvallis, Oregon, 20, 22, 23 July, 6, 23, 25, 30 August, and
4 September, also Forest Grove, 1 June; Blooming, 26 July; and Hills-
boro, 24 September, all in Oregon.

Type in Oregon State College, paratypes there and in the M. C. Z.
no. 25733.

Priocxemis nebulosus Dahlb.

Dahlbom says the wings are violaceous, mentions the emarginate
clypeus, and locality South Carolina. From Florida in the Graenicher
collection are two males and a female which agree better with the
description than the form from Virginia northward that, following
Cresson, I previously identified as nebulosus.

In the Florida female the clypeus is more plainly emarginate in the
middle, the angle each side is almost a tooth; the wings are more
evenly dark than the northern form, which will be known as P. pul-
chrina Cresson, based on the male.

In the male of nebulosus the wings are dark as in the female (in
pulchrina the male has hyaline wings with a smoky tip), and the black
median stripe on the face is broader, especially on the clypeus. The
legs are marked as in pulchrina. In both sexes the second submarginal
cell is not as long as in pulchrina.

A female from Larkins, May. males, South Miami, 26 April, and
Lake Apopka, Winter Garden, 25 April, all Florida.

Priophanes

There are several among the described species of Priocnemis which
have a distinct petiole connecting the abdomen to the propodeum,
while true Priocnemis has no petiole. For the petiolate forms I pro-
posed a new genus which is more related to Pseudagenia and Ageniella
than to Priocnemis. The venation is much like Pseudagenia, the basal
vein ends before the transverse, and in hind wings the anal vein ends
before the cubital fork. The hind tibiae have three rows of spines

174 bulletin: museum of comparative zoology

above the middle one with more or less distinct teeth; last joint of mid
and hind tarsi bare, the claws toothed; no distinct "beard" under
head; venter of female with a groove on second segment; the mesoster-
num not prominent laterally.

Type Priocnemis facetus Cress.

Also included are P. agcnoideus, arizonica, arcuatus, holonis, n. sp.,
placitus, relictus.

Priophanes holonis spec. nov.

Black; abdomen reddish all over; wings hyaline, apex broadly
dusky.

In general close to P. directa of Texas; the pronotum angulate be-
hind, the spurs black, legs and antennae entirely black; and head,
thorax, and abdomen of the same shape as P. directa. It differs in
having the teeth on hind tibiae much smaller, hardly noticeable, except
from inner side, and more especially in venation. The marginal cell is
longer, its outer side more oblique, and not quite its length from tip of
wing (in directa more than its length from tip of wing) ; the second and
third submarginal cells are proportionally longer than in directa, and
the lower outer corner of third submarginal cell is scarcely the length
of third submarginal from the outer margin of wing (in directa more
than length of third submarginal cell from outer margin of wing).
The first recurrent ends at or before middle of second submarginal,
and second recurrent before middle of third submarginal cell. The
basal vein, which is curved as in directa, ends only a little before the
transverse vein.

Length 7.5 mm. to 8.5 mm.

Holotype from Urbana, Illinois, 20 July (Bequaert); paratypes
Columbus, Ohio, August (Bequaert), also 6, 21 July (Gillaspy); and
MacCollum, Coweta Co., Georgia, 8 June (Bequaert). Type M. C. Z.
25892, paratypes there and Ohio State University.

Ageniella delicata spec. nov.

Dull black, abdomen with all of second, most of first, and part of
third segments yellowish rufous; tip of abdomen with white spot;
mandibles yellowish at tips; antennae brown, basal joint pale yellow
beneath; all coxae black, rest of front legs pale yellowish, mid legs
with tibiae and femora pale, tarsi brown, hind legs with femora pale,
tibiae and tarsi dark brown; spurs of hind legs dark brown, others
paler; wings hyaline, tip not darker, veins yellowish to brown.

banks: psammocharidae 175

( "lypeus and lower face with silvery pubescence; notum and pleura
somewhat sericeous.

Body slender; clypeus broad, somewhat rounded below; lateral
ocelli much nearer each other than to eyes; vertex convex; antennae
long and slender, third joint hardly longer than fourth; pronotum
deeply arcuate behind; propodeum finely granular, median groove
faint; basal segment of abdomen more than twice as long as broad
behind.

Fore wings rather short, the submarginal cells not as broad as the
marginal, latter nearly its length before tip of wing; second submargi-
nal about one-third longer than broad, receiving the first recurrent
vein near base; third submarginal plainly longer on lower side than
second, but narrowed nearly one-third above, receiving the second re-
current (almost angulated) a little before middle; lower part of basal
vein bulging forward, ending a little before the nervellus.

Mid and hind legs very slender and long, smooth, inner spur of hind
tibiae two-thirds of basitarsus.

Length of fore wing 4.2 mm.

One male from Falls Church, Va., 22 August. Type M. C. Z. No.
25910.

Ageniella restricta spec, now

Body dull black, more or less sericeous, most noticeable on the
coxae and pleura; clypeus and lower face with dense white pubescence;
abdomen with basal two-thirds of second segment yellowish rufous;
tip of abdomen with a white spot; legs brown to black, front legs with
tibiae and tarsi pale yellowish, hind spurs dark, others paler. Fore
wings dusky, tip rather darker, veins dark brown; hind wings almost
hyaline.

Body slender; clypeus truncate below; antennae slender, third joint
a little longer than the fourth, on the style of A. iridipcnnis; lateral
ocelli plainly nearer each other than to eyes; vertex rather strongly
convex; pronotum arcuate behind; propodeum finely granular, sloping
toward tip, median line scarcely visible; abdomen slender, basal seg-
ment more than twice as long as broad behind.

In fore wings the submarginal cells as broad as the marginal cell,
latter not quite its length from the tip of wing; second submarginal
nearly one and a half times as long as broad, receiving the first recur-
rent before basal third; third submarginal cell little longer below than
second, but broader, narrowed hardly one third above, receiving the
second recurrent (nearly evenly curved) before middle: lower part of

176 bulletin: museum of comparative zoology

basal vein only slightly bulging, and ending a little before the nervel-
lus; mid and hind legs long and smooth, inner spur of hind tibia more
than one-half but hardly two-thirds of basitarsus.

Length of fore wing 5 mm.

One male from Falls Church, Virginia, 22 August, on leaves of
tulip-tree with honey-dew. Type M. C. Z. No. 25909. Closely re-
lated to A. delicata, but differing in less reddish on abdomen, basal
joint of antennae black, shorter spurs, darker wings and venation, and
broader submarginal cells.

Ageniella neglecta spec. nov.

Body, antennae, legs, wholly black, hind spurs black, others paler;
fore wings nearly hyaline, slightly darker near tip, veins brown.
Clypeus and lower face with white pubescence, thorax and coxae
somewhat sericeous, not on abdomen.

Clypeus truncate below; antennae short and rather thick, the third
joint little more than twice as long as broad at tip ; pronotum broadly
arcuate behind; propodeum (from side) slightly rounded, no median
line; basal segment of abdomen not quite twice as long as broad be-
hind. Legs short, smooth, inner spur of hind tibiae little more than
one-half of basitarsus.

In fore wings the marginal cell is fully its length before tip of wings,
the third submarginal cell nearly three times its length from margin,
and almost as broad as the marginal cell; second submarginal cell a
little longer than broad, receiving the first recurrent at basal third;
third submarginal a little longer than the second, only a little nar-
rowed above, receiving the slightly curved second recurrent near tip;
lower part of basal vein only a little convex and ends on the nervellus ;
in hind wings the anal ends much before cubital fork.

Length of fore wing 3.5 mm.

One male from Boulder, Colorado, 26 August, 1908 (S. A. Rohwer).
Type M. C. Z. No. 25908.

Readily known by small size, deep black color, short legs, and the
large apical field beyond venation.

Ageniella accepta var. conflicta var. nov.

Differs from the typical form in having a black spot on side of scutel-
lum and on side of metanotum as well as at base of fore wing, usually
all run together in a black stripe; area between ocelli black; body more

banks: psammochajridae 177

slender than typical form; the pale area of fore wing between the first
brown band and the stigma] band is more suffused with pale brown, so
that the wing is not so plainly three-banded; the basal part of wing
is more yellowish, and the brown at apex is a paler brown than in
typical acccpta.

Holotype, a female from Falls Church, Va., 5 July; paratypes also
from Falls Church from 19 June to 13 Sept.; from Glencarlyn, Va.,
26 July, and from Riverhead, L. I., X. Y., 1 August (W. T. Davis).
Type M. C. Z. no. 25735.

Synoptic table of males of eastern species
of Ageniella at present known to me

1. Tip of abdomen with a distinct white spot; third cubital cell longer below

than above 2

Tip of abdomen wholly dark 11

2. Some reddish or yellowish on dorsum of abdomen; basal segment of abdo-

men more than twice as long as broad 3

No reddish nor yellowish on dorsum of abdomen 7

3. All spurs snow-white ; femora rufous birkmanni

At least hind spurs dark 4

4. Clypeus with a small pale spot on each side, and a small orbital spot .festina-
Clypeus wholly dark 5

5. A small yellow orbital line or spot; marginal cell much less than its length

from tip of wing; inner spur of hind tibiae scarcely more than one-half of

basitarsus; basal joint of antennae dark fratemella

Marginal cell nearly its length from tip of wing; no pale orbital line or spot;
inner spur of hind tibiae more than one-half of basitarsus 6

6. Basal joint of antennae pale yellowish below, femora yellowish or rufous,

paler than rest of legs; red of abdomen on all of second and parts of first

and third segments delicata

Basal joint of antennae black below; femora more brown, darker than rest
of legs; red on abdomen only on part of second segment restricta

7. Two pale spots or a white band on hind border of pronotum ; clypeus mostly

whitish; spurs snow-white calcarata

No pale mark on pronotum 8

8. A small yellowish spot or short stripe on orbital line a little above clypeus;
long spur of hind tibiae about one-half of basitarsus; vertex usually shin-
ing; fore wings more than five millimeters long agilis

No such spot or stripe on orbital line; fore wings not over five millimeters
long 9

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9. Femora rufous or yellowish, sometimes tibiae also; inner spur of hind

tibiae hardly more than half of basitarsus texana

Femora and tibiae black; inner spur of hind tibiae more than one-half of
basitarsus 10

10. Abdomen very slender, second and third segments with pendent side-mem-

brane, basal segment plainly more than twice as long as broad behind.

pctiolata

Abdomen broader, no side-membrane to second and third segments; basal

segment twice as long as broad behind, tip of fore wing black norata

11. Abdomen with some reddish or yellowish on basal half above 12

Abdomen wholly black above 14

12. Basal segment of abdomen not twice as long as broad behind; third sub-

marginal cell with angulate hind border perfecia

Basal segment of abdomen more than twice as long as broad; small species.

13. Clypeus black; second segment of abdomen dark in middle, yellowish on

sides; third submarginal cell higher than long; hind spurs dark; basal
segment of antennae black; third submarginal angulate behind.

minuscula

Clypeus mostly pale; spurs white; basal joint of antennae pale above and

below; hind border of third submarginal cell not at all angulate behind.

apicipennis

14. Clypeus with some pale, at least on lower edge; palpi pale 15

Clypeus wholly dark 17

15. Third submarginal cell with outer border angled in middle, so cell no

broader below than above; front coxae almost white clypeata

Third submarginal cell plainly longer on lower side than on upper 16

16. Hind border of pronotum slightly pale; basal joint of antennae white below.

hestia
Hind border of pronotum shows no paler band; basal segment of abdomen
not twice as long as broad behind; antennae thicker than usual.

crassicornis

17. Third submarginal cell with hind border angled in middle, so cell no

broader below than above 18

Third submarginal cell with outer side sloping, and lower border longer
than upper 19

18. Hind spurs dark; second recurrent arises near the middle of outer part of

cubitus; small species eximia

All spurs nearly white; second recurrent arises plainly beyond the middle of
outer part of cubitus ; larger species osoria

19. All spurs snow-white; basal segment of abdomen fully twice as long as

broad behind virginica

Basal segment not quite twice as long as broad behind; all spurs not white

banks: psammocharidae 17!)

20. Third suhmarginal cell much longer than high; a distinct median groove

on propodeum species?

Third submarginal cell scarcely as long as high, often plainly higher than
long below; no distinct groove on propodeum 21

21. Hind femora reddish (or yellowish) on outer half, base black or brown.

aludra
If hind femora pale it is also on basal part 22

22. Second recurrent arises little if any beyond middle of outer part of cubitus.

23
Second recurrent arises plainly beyond middle of outer part of cubitus . . 24

23. Front femora, tibiae, and tips of coxae pale; third submarginal cell always

longer below than above; hind legs often partly pale tenella

Front femora, tibiae, and coxae not pale; third submarginal cell often no
longer below than above; legs black, a small species eximia

24. Front tarsi yellowish; basal segment of abdomen about one and one-half

times as long as broad behind iridipennis

Front tarsi not yellowish; basal segment of abdomen not one and one-half
times as long as broad behind atrata

Nemagenia subgenus nov. of Ageniella

Pronotum above as long as mesonotum ; marginal cell its length from
tip; basal vein plainly before nervellus; inner spur of hind tibiae not
one-half of basitarsus; propodeum very long, nearly flat, from side
it is only slightly curved; otherwise like Ageniella.

Type Pompilus (Agenda) longulus Cresson.

Only the male is known; described from North Dakota, I have
specimens from Fedor, Texas (Birkmann).

Dipogon texanus spec. nov.

Head and abdomen black, clypeus and thorax rufous, legs dull
black to brown, tarsi partly paler, coxae rufous, antennae yellowish,
a narrow-black band at tip of each joint, except basal and apical; fore
wings hyaline, a narrow black band across over the basal vein, and a
very broad blackish band across over submarginals and third discoidal
cells, not occupying the tip of marginal cell.

Face and vertex with short appressed white hair ; abdomen also with
pale appressed hair, longer and erect hair near tip; venter with bands
of pale hair. Hair basket under head of white bristles. Structure in
general similar to D. brevis; very little hair on thorax or propodeum;
venation similar, but marginal cell hardly as broad, but angulate on
hind border; second submarginal about twice as long as the third; first

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recurrent ending much before middle of second submarginal, second
recurrent ending near base of third submarginal cell; medius reaches
margin; legs slender as usual; long spur of hind tibia about two-fifths
of basitarsus.

Length 6.5 mm.

From Brownsville, Texas, 11 to 16 June (Darlington). Type
M. C. Z. No. 25896.

Dipogon sericea spec. nov.

Body black; antennae with first and second joints black, beyond
yellowish, some joints narrowly dark at tips; legs black, tarsi and the
front tibiae rufous. Fore wings brown, beyond the marginal cell
snow-white, the marginal cell and below and beyond it darker than
elsewhere, stigma black; hind wings slightly infuseate.

Thorax and abdomen densely clothed with appressed gray pu-
bescence, in places somewhat yellowish, thorax with rather long, erect
white hairs, and shorter ones on the abdomen, near tip darker. Face
above antennae with bright yellowish pubescence, across face half way
up to vertex is a band of erect black bristles, and similar bristles on
vertex; the lower edge of clypeus is yellow, rest black, clothed with
white hairs, white hairs back of eyes; hair basket of fine, pale bristles.

Legs with the femora, mid and hind tibiae covered with sericeous
pile, the femora with fine, long hairs below. Hind tibiae above with a
groove and a row of fine short hairs, long spur about one-third of the
basitarsus. Antennae rather slender, third joint not much longer than
fourth, together they equal vertex-width.

Thorax shorter than in Eastern species, but fully as broad ; pronotum
broadly arcuate behind; propodeum appears to have a broad median
furrow, but all covered with the appressed hair.

In fore wings the venation is much like the Eastern D. sayi except
that the marginal cell is much shorter, extending only a trifle beyond
the third submarginal cell; the medius reaches to the margin, but in
the white area it is very fine. In hind wings venation as in other
species.

Length of fore wing 6 mm., body 6.5 mm.

One from Bull Prairie, Lake Co., Oregon, 22 July, Camas Prairie
Summit, 7,500 ft. (Frewing coll.) Type at Oregon State College.

The species of Dipogon so far described from the United States
can be separated by the following key :

1. The medius of fore wing plainly does not reach the margin; marginal cell not
strongly angled at end of second submarginal cell, the outer side curved.

Subgenus Adipogon — 2

banks: psammocharidae 181

The medius reaches to the outer margin; the marginal cell is strongly
angled at the end of the second submarginal cell, and broadest at this point.

Subgenus Dipoyon — 3

2. Forewings wholly dark, nearly black papaya

Forewings clear with a narrow dark band over the basal and transverse

veins, and a large dark spot over marginal cell and the two or more cells
behind it pulchripennis

3. Thorax yellowish to rufous 4

Thorax black 5

4. Head and abdomen black; antennae rufous, tips of joints narrowly black;

legs partly black texan us

Head rufous, abdomen black; antennae rufous, no dark bands; legs wholly
yellowish yraenicheri

5. Face, mesonotum, and abdomen with appressed grayish to yellowish pu-

bescence; hair basket pale; front legs partly pale 6

Face, thorax, and abdomen without such pubescence; fore wings clear with
two bands 7

6. Across face and on vertex are erect black bristles; fore wings brown, tip

snow-white; propodeum and pleura also sericeous sericea

No such bristles on face and vertex; extreme wing-tip faintly dark, no snow-
white; propodeum and pleura without appressed hair brevis

7. Front legs largely yellowish, also mandibles, and lower edge of clypeus.

caliptera
Front legs, clypeus, and mandibles black sayi

Adipogon, subgenus nov. has the basket of curved hairs on under
side of head as in typical Dipogon, but the median vein of fore wing
does not extend to the outer margin, and the marginal cell is not
angled below.

Type is Pompilus pulchripennis Cresson.

Pepsis pattoni spec. nov.

Our large black Arizona Pepsis has been identified by Fox and my-
self as P. obliquerugosa Lucas, a form described from Cuba. A few
years ago Salman considered both to be P. grossa Fabr. of northern
South America. With a series of both sexes of the three forms I con-
sider each a separate species.

As with the others, the wings are black with a narrow pale band at
tip, not as broad, and in the female not as white as in obliquerugosa.
On the propodeum the side tubercles are very prominent, sometimes
almost pointed, in obliquerugosa much lower and less noticeable. At

182 bulletin: museum of comparative zoology

the middle of the turn there is in both species a high ridge; in obliqueru-
gosa its top is straight or a little convex, in pattoni the top is plainly
emarginate in the middle. The oblique ridges on the propodeum so
characteristic of obliqucrugosa, are fewer, less oblique, and sometimes
none oblique. In pattoni the third joint of antennae in female is a
little shorter than in obliqucrugosa in specimens of same wing-length.
In the male the parameres of the genitalia are slender and tapering
toward tip; in grossa (see Lucas figure 24) they are heavier, and not at
all tapering; in obliqucrugosa the parameres are not as broad as in
grossa, but do not taper as much as in pattoni; in the latter the tip of
the subgenital plate is broadly convex, in the Cuban form it is plainly
emarginate in middle. The females of all three forms have long, stiff,
curved bristles under the front femora, and with our ncphclc and a few
South American species they form a natural group. Lucas placed the
male of ncphclc asformosa and says that the genitalia are the same as in
grossa, and gives no separate figure. The males of this section have
the subgenital plate slender and furnished with a broad dense crest of
long black hairs.

P. obliqucrugosa is somewhat the largest of the three species, fore
wing sometimes 45 mm. P. pattoni sometimes about 42 mm., fre-
quently 40 mm. P. grossa rarely 40 mm., often near 35 mm.

Holotype of P. pattoni 9 is from Palmerlee, Arizona, July (Bieder-
mann coll.), M. C. Z. No. 25805; allotype from southern Arizona;
paratypes are from Palmerlee, Tucson, Ola, Ft. Grant, Pinalerro
Mts., Santa Catalina Mts., all Arizona, from late June to September,
taken by Snow, Bequaert, Morse, and Wheeler. One from mountains
near Pomona, California, by H. C. Fall.

b. ANTILLEAN

Batazonus gundlachi Cress.

Described from female only.

Male. It is very similar to female, and marked the same; thorax
and abdomen brownish red, the clypeus is reddish in middle, above
antennae are two short curved dark streaks, there is some yellow each
side of scutellum (as well as the post-scutellum) ; a large yellow spot
just above base of mid coxae; propodeum broadly yellow across apical
half; abdomen black at extreme base, and faintly dark across tip of
first segment, a broad yellow band across base of third segment above
and below; legs reddish, except the paler tarsi which are dark at tips

banks: psa mmoch arid ae 183

of joints; inner spur of hind tibia about three-fifths of basitarsus. The
third submarginal cell is much longer than high, as in the female.

Length of fore wing 13 mm.

Allotype from San Bias, Trinidad Mts., Cuba, 24 April (G. E. Folk).

Batazonus hookeri Rohwer

Described from female only.

The male is largely yellowish; there is a triangular black spot below
antennae, two broad black stripes above, narrowly separated; two
yellow lines on mesonotum, the mesopleura has two slightly separated
elongate, yellow spots; the propodeum above is yellow, black at ex-
treme base, usually extended back in middle to divide or partly divide
the yellow. The dark bands on abdomen above are brown to black,
occupying hardly one-half of the segment, basal segment dark only at
tip; coxae black, with a yellow mark; femora partly dark, tip pale; tips
of tarsal joints black; inner spur of hind tibiae about four-fifths of the
basitarsus.

Length of fore wing 5 to 7 mm.

Allotype from Hatillo, Puerto Rico, January ; others from Mayaguez
in November and Cartagena in May, both Puerto Rico, all through
Mr. Ramos.

PSAMMOCHARES PARSONSI Spec. nOV.

Body, legs, and antennae wholly deep black; wings mostly black, but
paler in third discoidal cell and toward base both in front and behind;
hind wings strongly fumose.

Clypeus over three times as broad as long, nearly truncate below,
a few hairs on lower part; ocelli in a low triangle, the laterals a little
nearer to the eyes than to each other; a median line above base of
antennae, latter slender, third joint equal vertex-width, last joint only
about one-half of third joint; vertex with a few long, erect hairs,
shorter black hair on front.

Pronotum broadly arcuate behind; mesonotum with a few long hairs
each side, pleura with only scattered short hairs; propodeum short, no
distinct median groove, with rather short hair above.

Abdomen hairy toward tip below, above with stiff bristles just be-
fore tip, otherwise a few hairs on venter near hind border of segments.

Femora smooth, without hairs; front tarsi with a distinct, though
short comb, a spine at middle of the second tarsal joint; hind tibiae
quite heavily spined; two irregular rows above, some as long as the

184 bulletin: museum of comparative zoology

width of the joint; inner spur of hind tibia more than one-half of
basitarsus.

In fore wings the marginal cell is about its length before tip ; second
submarginal cell longer below than high, one-third narrowed above,
receiving the first recurrent vein near tip; third submarginal cell only
a little longer than second and likewise narrowed one-third above,
receiving the second recurrent vein (evenly curved) beyond the
middle. In hind wings the anal ends beyond the fork.

Length of fore wing 10 mm.

Females from Buenos Aires, Trinidad Mts., Cuba, 17 to 23 June
(C. T. Parsons).

In appearance like P. pcrpilosus but easily separated by absence of
hairs on femora and very much less hair on body. Type M. C. Z.
No. 25742.

NOTIOCHARES ANTILLANA spec. nov.

Male. Very much like AT. cvbensis, wholly black, with blue and
violet reflections, and agreeing closely in structure to that species.
The only difference of importance is that the male genital plate, which
is N . cubensis ends in a sharp-pointed tooth each side, in N. antiUana
ends in short, rounded lobe, alike in all three specimens.

Length of fore wing 8 to 11 mm.

From Barbados, August to December (Spencers). Type M. C. Z.
No. 25743.

Episyron cressoni Dewitz

The male is black; a white spot on each side of clypeus; a broad
orbital streak yellowish white, reaching nearly to vertex; a narrow
yellowish band across tip of pronotum, a white spot each side on basal
part of third abdominal segment, the last dorsal segment somewhat
rufous, but white at tip; the mid femora rufous except base and tip,
the hind femora rufous except basal third, and the hind tibiae rufous
except extreme tip, spurs are pale; inner one of hind tibia nearly equal
to basitarsus.

Length of fore wing 6 mm.

Allotype from Mona Island, April (Ramos).

Priocnemis ursula spec. nov.

Body black, much of it covered with a fine sericeous pile or bloom;
clypeus, mandibles, basal joint of antennae, and a few joints beyond

banks: psammocharidae IS.")

yellowish, rest of antennae brown; legs black; wings hyaline, venation
black.

Clypeus above about as broad as vertex, nearly evenly rounded
below, fully two and one-half times as broad as long, with a few fine
hairs; front with fine, pale hairs, and vertex with some longer ones;
ocelli in nearly equilateral triangle, laterals a little (but not much)
nearer to each other than to the eyes; a short median groove above
base of antennae, latter slender, second plus third joints not as long
as vertex-width; pronotum scarcely constricted behind, with a broadly
arcuate hind margin ; mesonotum not humped in middle, scarcely hairy,
scutellum and metanotal lobe with erect black hair; propodeum with
rather long, fine, pale hairs above.

Abdomen hairy only toward tip and somewhat beneath; legs
moderately slender, hind tibia above with about ten teeth with a
short spine, and a row of spines on outer part; inner spur of hind tibiae
two-fifths of basitarsus.

The fore wings have the marginal cell rather longer than space to the
tip of wing, three times as long as broad, outer side nearly straight;
second submarginal cell oblique, below nearly twice as long as high,
receiving the first recurrent vein beyond the middle; the third sub-
marginal cell plainly longer than second, but not as much wider as in
many species, outer margin bent near median vein, receiving the
second recurrent vein (almost straight) much before middle; in hind
wings the anal vein ends much before the fork.

Length of fore wing 6.5 mm.

Female from Villa Altagracia, San Domingo, July (Darlington)
Type M. C. Z. No. 25741.

In general very similar to P. salti Bks. which differs in the yellowish
tip to the abdomen.

Priocnemis arioles spec. nov.

Head and thorax black, abdomen reddish except black tip, the fifth
segment above bluish, thoracic notum bluish, legs black, femora and
tibiae somewhat bluish; antennae black; fore wings nearly evenly
infumate, slightly darker toward costal tip than behind, hind wings
slightly smoky.

Clypeus no wider than face below, with a few pale hairs, about two
and one-half times as broad as long, truncate below; vertex as broad
as face below, scarcely convex, with a few erect hairs each side, none
on front; ocelli in a rather low triangle, the laterals much nearer each

186 bulletin: museum of comparative zoology

other than to eyes ; a groove from anterior ocellus to base of antennae.

Pronotum plainly contracted behind, the margin slightly arcuate;
mesonotum almost humped in middle, with one or two erect bristles
each side and a pair on scutellum, pleura bare; propodeum with only
faint, erect, pale hair above, a whitish pile across base, and an elongate
spot each side toward hind margin, surface minutely granulate, no
median groove. Abdomen hairy on venter and apical third above.
Legs not especially slender, mid tibiae with two rows of short spines
above, hind tibiae with a row of seven or eight oblique teeth, a spine
beyond each, and an outer row of short spines ; inner spur of hind tibia
hardly two-fifths of basitarsus.

Marginal cell of fore wings about its length from wing-tip, two and
one-half times as long as broad, outer side convex; second submarginal
cell a little longer than high, but little narrowed above, receiving the
first recurrent vein at middle ; third submarginal cell much longer than
second and much wider behind, one-third narrowed above, receiving
the second recurrent vein (slightly curved) a little before middle; in
hind wings the anal vein ends much before the fork.

Length of fore wing 6 mm.

Female from Constanza, Valle Nuevo, San Domingo, 21 August,
3- to 4000 ft. (Darlington). Type M. C. Z. No. 25740.

The Antillean species with a more or less reddish abdomen can be
tabulated as follows : —

1. Fore wings hyaline with two broad dark bands pulchellus

Fore wings without bands 2

2. Thorax wholly reddish christophei

Thorax black 3

3. Third submarginal cell no longer than second, higher than long parous

Third submarginal cell much longer than second, and much longer than

high 4

4. Wings hyaline; hind legs very long, body about 4.5 mm dowi

Wings plainly infuscate, hind legs not so long, body 7 mm ariolcs

Priocnemella domingensis spec. nov.

Head, propodeum, and pleura black, thorax and abdomen above
blue. Fore wings black, violaceous, hind wings infuscated, also with
violet reflections; antennae and legs black, the femora and tibiae
above, and the front coxae iridescent bluish, also basal joint of anten-
nae above.

Clypeus large, extending laterally under the eyes, with long, black

banks: psammocharidae 187

hairs, lower margin narrowly smooth and coming to a blunt point in
middle; front and vertex with long black hair, one each side on vertex
very long; third antennal joint equal to vertex-width. Ocelli close to-
gether, the laterals more than twice as close to each other as to the
eyes; groove from anterior ocellus to the antennae. Scattered erect
black hairs on notum and scutellum, short hair on pleura; pronotum
almost angled behind. Propodeum minutely, transversely striate,
more distinctly so than in P. violaceipes, above and on sides with
sparse, erect black hair.

Abdomen hairy near tip and below, scarcely so near base. Legs
slender, spined as in P. violaceipes, the mid and hind tibiae with two
rows of erect, short black spines, no teeth, and evenly short spines on
the tarsal joints; claws with an erect, small tooth; long spur of hind
tibia about two-fifths of the basitarsus.

Venation of wings as in P. violaceipes; the marginal cell long and
pointed; second submarginal oblique, about one and one-half times as
long as high, receiving the first recurrent vein near middle; the third
submarginal cell much longer, about one-third narrowed above, re-
ceiving the second recurrent also near middle, this recurrent is more
sinuously curved than that in P. violaceipes.

Length of fore wing 10.5 to 11 mm.

Two females from near and southeast of Constanza, Valle Nuevo,
San Domingo, August, 3 to 7000 ft. (Darlington). Type M. C. Z. No.
25739.

Except for the striking difference in color of thorax and abdomen,
and slight differences in structure, it is practically the same as P. vio-
laceipes of Cuba.

Priochilus

A few Neotropical species have much the appearance of Priocnemis;
however, the last joint of hind tarsi has spines beneath, but not later-
ally; the hind tibia has no real teeth like Priocnemis, but spines in
rows, •some as long as the diameter of the joint ; the claws are cleft, the
inner part broader than the outer part, and somewhat obliquely trun-
cate; the palpi are slender as in Priocnemis.

Pompilus nobil is Fabr. is the genotype; Salius opacifrons Fox from
Jamaica goes in the genus, although smaller than most others. In
South America there are regius Fabr., diversus Smith, scrupulus Fox,
sericeifrons Fox and others. Mr. Williams in his paper "Studies in
Tropical Wasps", 192S, p. 141, calls attention to nobilis and regius as
possibly forming a new genus.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 5

SCIENTIFIC RESULTS OF A FOURTH EXPEDITION
TO FORESTED AREAS IN EAST AND CENTRAL AFRICA

VI
ITINERARY AND COMMENTS

By Arthur Loveridge

With Four Plates

CAMBRIDGE, MASS., U.S.A.

PRINTED FOR THE MUSEUM

July, 1944

PUBLICATIONS
OF THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
investigations by the Staff of the Museum or of reports by spec-
ialists upon the Museum collections or explorations.

Of the Bulletin, Vols. I to XCIII, and Vol. CXIV, No. 1, 2,
3 and 4 have appeared and of the Memoirs, Vol. I to LVI.

After 1941 no more Memoirs are to be published.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 5

SCIENTIFIC RESULTS OF A FOURTH EXPEDITION
TO FORESTED AREAS IN EAST AND CENTRAL AFRICA

ITINERARY AND COMMENTS

By Arthur Loveridge

With Four Plates

CAMBRIDGE, MASS., U.S.A.

PRINTED FOR THE MUSEUM

July, 1944

No. 5. — Scientific Results of a Fourth Expedition to Forested Areas in

East and Central Africa

Itinerary and Comments

By Arthur Loveridge

INTRODUCTION

For various reasons it would appear inopportune at the present
juncture to devote the time necessary to elaborating the ecological and
zoogeographical data accumulating from this, and previous, expedi-
tions so generously sponsored by the John Simon Guggenheim Me-
morial Foundation of New York.

On several occasions, however, correspondents in Africa have
written requesting specific information about, or the latitude and
longitude of, some of the more obscure localities mentioned in the
reports already printed, so that it would seem advisable to publish
the itinerary without further delay. A synopsis of it has, indeed, been
furnished in the caption accompanying Plate 1, and it has not been
thought necessary to enlarge on this in respect to places where only
a single night was spent, with consequently little collecting.

For the others I am now supplying the latitude, longitude, altitude,
the all-important meteorological conditions prevailing at the time of
my visit, duration of stay, and position of camp. This in turn is fol-
lowed by a brief survey of the vegetational environment, or a reference
to where such an account has been published elsewhere.

The composition of the fauna in relation to life zones receives atten-
tion, more particularly for localities in Tanganyika Territory where it
has greater significance on account of our knowledge being more com-
plete. When one contemplates the vastness of the largely trackless
forests in Uganda, one feels that the results accruing from a stay of
two weeks in areas of from 120 to ISO square miles in extent, is alto-
gether too inadequate to justify one in embarking on detailed specu-
lation as to what does, or does not, occur.

Species of exceptional interest receive mention where it is thought
that they may assist in visualizing the environment, or where, because
of their rarity, other zoologists visiting the locality would wish to
know of their occurrence. Special reference is made to forms of which
the type happens to have come from the particular locality under
discussion.

192

bulletin: museum of comparative zoology

The groups already reported upon in this Bulletin are as follows :

Speci-

New to

Volume

Pages

Class

mens

Species

M.C.Z.

145-214

I Mammals

812

116

215-275

II Birds

809

240

183-234

III Crustacea

389

235-374

IV Reptiles

1802

151

375-436

V Amphibians

1081

It has been somewhat of a disappointment that no one has been able
to undertake the identification of, or to report upon, the fairly exten-
sive collections of other invertebrate groups such as mollusks, myria-
pods, and earthworms. In addition to zoological material, we pur-
chased from Baamba and Banyaruanda tribes 1013 ethnological
items, with a gross weight of almost one ton, which were sent by
truck and train over a thousand miles to the port of shipment.

For part of the data regarding altitudes, temperature, composition
of forests, etc., I am indebted to "Uganda" by Messrs H. B. Thomas
and R. Scott, 1935, and to the "Handbook of Tanganyika Territory"
by G. F. Sayers, 1930; both mines of reliable information.

There remains only to again express my deep gratitude to the John
Simon Guggenheim Memorial Foundation, and to Dr. Thomas Bar-
bour, Director of the Museum of Comparative Zoology, for making
possible this expedition, of whose harvest of herpetological material I
am now able to take full advantage in the revisionary studies of
African lizards upon which I am engaged.

ITINERARY
UGANDA

Mabira Forest, Kyagwe (Chagwe). 0°24' N., 33°0' E. Alt. 4000 feet.

As the rains were still continuing we accepted the generous offer of
Mr. L. Jarvis to occupy the late Major Cuthbert Christy's old house
at Mubango, a rubber and coffee plantation in the southern end of
the forest, five miles north from Najembe, which is twenty miles
southeast of Jinja on the Jinja-Kampala road.

Arrived at noon on November 5, and left on the 21st.

loveridge: African itinerary and comments 193

A heavy shower amounting to .76 inches fell on the afternoon of our
arrival, .17 the following day, and .75 on the 14th. Actually a shower
or two occurred almost daily but the precipitation was too small to
be registered. The average annual rainfall at Mubango is 55 inches.
The daily temperature at 7 a.m. averaged 66°, and at noon 80°.

We had come here largely because Major Christy's labours had made
the Mabira, which covers 120 square miles, type locality for Leptopelis
n. christyi, Hylambates verrucosus, Miodon g. christyi, and Aparallactus
christyi (= A. modestus). Of these we were successful in securing only
the first and last.

Despite what appeared to me to be ideal conditions, I personally
captured only three snakes during as many days primarily devoted to
searching for them both in the forest and along its edge, where drifts
of leaves between buttress roots and rotting logs provided suitable
retreats. On four succeeding days I employed an allegedly expert
snake-catcher who got nothing during that entire period. Yet some
two-score labourers, engaged in clearing undergrowth on the planta-
tion, brought in snakes at the rate of nearly two a day, and of these
the first eleven snakes represented ten different species ! This astonish-
ing variety continued until we actually had eighteen species for a total
of twenty-seven snakes. Of these one — Dipsadoboa unicolor — was new
for Uganda, another — Bothrophthalmus I. lineatus — constituted the
most easterly record for the species, though its presence here had been
forecast by Lt. Col. Pitman.

Pitman (1934, Uganda Journal, 1, pp. 7-16) has furnished an inter-
esting account of the Mabira and its fauna and flora, rendering un-
necessary further details here. Lizards, with the exception of the
arboreal Algiroides africanus, were disappointing, being of widespread
species. A single chelonian was encountered.

The dense undergrowth rendered bird collecting extremely difficult
and only twenty skins were prepared during the fortnight. More than
a score of mammals were taken, however, including such forest forms
as Cercocebus, Cercopithecus, Hcliosciurus, Tamiscus, Protoxerus
Cephalophus and an arboreal pangolin which was described as a new
race (Phataginus tricuspis mabirae).

Budongo Forest, Bunyoro. 1°42' N., 31°24' E. Alt. 4000 feet.

Camp was pitched in the mahogany nursery at BISU beside the
Buchanan Saw Mills, so that full advantage might be taken of the
three-mile-long corridor cut into the heart of the forest.

Arrived in afternoon of November 22, and left on December 7th.

Heavy downpours occurred at intervals during our stay for the rain-

194 bulletin: museum of comparative zoology

fall here is well distributed throughout the year, the annual average
being 65 inches. During our visit the daily temperature at 7 a.m. aver-
aged 60°, and at noon 82°.

Situated on the north-north-westerly slope above the escarpment
towards the northern end of Lake Albert, Budongo is rich in mahogany
and consequently considered the most valuable of all Uganda's lovely
forests. Ironwood trees are also plentiful, in some areas forming over
50% of the canopy and being largely responsible for the maintenance
of its evergreen appearance. The forest survey reveals that the second
story stratum, attaining about 80 feet, is composed in part of such
genera as Celtis and Funtumia spp., while the third story is scanty.
Bordering the forest and at the edge of clearings, a dense, often im-
penetrable, undergrowth of bush has sprung up; but in the depths of
the forest where the canopy was contiguous one might wander at will
or chase the frogs which went leaping over the damp leaves that car-
peted the forest floor.

Our visit added four amphibia to the Uganda list, the most in-
teresting being Rana christyi, for Budongo, covering about 180 square
miles, shows very close faunal relationship with the great Ituri Forest
lying away to the west in the Belgian Congo.

This is borne out by a number of interesting herpetological records
recently resulting from the collections made here by Mr. W. J. Eggel-
ing, Conservator of Forests, whom we had the pleasure of meeting.
This West African complexion was reflected by 7 of the 9 species of
snakes, and 4 of the 7 kinds of lizards collected, though one of these —
Algircrides africanus, is more central.

Budongo is type locality for 7 valid, and 2 invalid, races of birds,
only one of which I succeeded in getting, this was the gorgeous little
forest kingfisher {Myioceyx lecontei ugandae) which obligingly alighted
on a stump in camp.

Of the 10 species of mammals secured, only 4 can be considered
definitely western, the best being a huge horseshoe bat (Hipposideros
cyclops) netted in a clearing deep in the forest, where was also the great
tree on which I shot four small chipmunk-like squirrels (Tamiscus
alexandri). Here too we trapped a strange harsh-furred mouse (Lop-
huromys aquilis subsp.) whose race proved unidentifiable; perhaps a
series might reveal it as an undescribed subspecies.

Kibale Forest, Tow. 0°24' N., 30°24' E. Alt. 4200 feet.

Camp was made near the Duta River where it crosses the new road
being cut through the forest about 20 miles southeast of Fort Portal.

Arrived late on December 8, and left on the 18th.

loveridge: African itinerary and comments 195

Thunderstorms, accompanied by rain, occurred almost every after-
noon, while on two days an exceptionally heavy downpour continued
on throughout the night. The annual average rainfall is 56.87 inches.
During our stay the daily temperature at 7 a.m. averaged 59°, and at
noon 77°.

Kibale, together with the adjacent Itwara and Muhangi Forests,
covers about 276 square miles. The northern portion, traversed by
the Kampala-Fort Portal Road, is impenetrable on account of the
dense thicket undergrowth. Acting on the advice of Mr. W. J. Eggel-
ing, therefore, we selected the southern portion as being more open,
the enormous buttressed trees sufficiently dense as to prohibit under-
growth in many areas. After penetrating the forest about a mile-and-
a-half Ave had to camp for the Dura River was still unbridged; beyond
the river the road continued for another two-and-a-half miles through
beautiful forest, and it was along this stretch that most of my collecting
was done.

Excursions along this road during and immediately after heavy rain
were productive of sundry snails, slugs and worms, the latter very
numerous and one species frequently a couple of feet in length. This
larger species, with a diameter as great as one's thumb, was remarkably
iridescent. Beyond the forest proper these worms were encountered
in muddy soil beside a stream which flowed through a patch of palm
forest much frequented by elephant. Butterflies in bewildering variety
surpassed anything which I had seen anywhere in East Africa.

It was disappointing, therefore, to find the herpetofauna extremely
scarce, not merely in species but in individuals also. All our efforts re-
sulted in obtaining only 7 species of reptiles and 7 of amphibians,
though there was some compensation in the fact that all were exclu-
sively sylvicoline species with the exception of Agama atricollis and
Bujo r. regularis which are equally at home in the savanna. The head-
man of the gang engaged in felling trees with which to bridge the
Dura, informed me that they encountered from 1 to 3 snakes only per
month. During the week the only one they brought me was a Rham-
nophis a. elgonensis. It was this paucity of poikilothermous creatures
that decided me to leave after ten days instead of remaining three
weeks as originally planned.

Monkeys were the dominant form of animal life and no fewer than 5
species, of which I collected 4, were observed feeding within a hundred
yards of my tent, the source of attraction being the huge spherical fruit
borne by a certain tree. Of a dozen species of mammals collected, a
squirrel (Funisciurus p. victoriae) was the only novelty.

196 bulletin: museum of comparative zoology

Bundibugyo, Tow. 0°41' N., 29°56' E. Alt. 3200 feet.

Stayed at the comfortable rest camp at Saza headquarters for
Bwamba District.

From the evening of December 19 to 26th.

A few showers occurred during the week. No annual rainfall record
was available as the area has only been opened up recently, but statistics
are being collected at the mission. During our stay the daily tempera-
ture at 7 a.m. averaged 55°, and at noon 62°.

This area, marked on the Uganda Survey Map of 1928 as Bwamba
Forest, is now largely covered by native gardens, including plots of
coffee, cotton, and bananas, with extensive rice cultivation in the
swampy areas. Elsewhere rank grass fifteen feet high, scattered trees,
clumps of indigenous oil palms (Elacis guinecnsis) and patches of
forest. The latter, said to amount to 140 square miles, is an extension
of the great Ituri Forest on the opposite bank of the Semliki River.

Naturally, therefore, sylvicoline snakes are plentiful and a large col-
lection could have been made had we been able to stay longer. At
first, however, the Baamba mutilated them so badly that many had to
be rejected and it was only during the last few days that they began
to arrive in good condition. At least 14 of the 16 species secured were
forest forms, of which Boiga puherulenta and Pscudohaje goldii were the
second records for Uganda and Miodon g. collaris the first for this race.

Three of the five kinds of lizards taken were widespread savanna or
eastern forest forms. No attempt was made to collect birds and the
only ones preserved were the swamp-loving rail (Sarothrura p: cen-
tralis) and warbler (Prima m. immutabilis) .

Except for a young duiker, all 42 of the mammals preserved were
rodents representing 9 species which reflected our position in the west-
ern foothills of the Ruwenzori Mountains overlooking the Semliki
River, some being montane species, others races of savanna rats.

The Baamba, who give their name to this region, are a forest tribe
of semi-pigmy stock who would undoubtedly prove helpful to a natural-
ist working in this country; unfortunately, as already indicated, de-
forestation has resulted in encroachment by many savanna forms
which would probably preponderate among specimens brought in.
The big tree cobra was obtained by Bakonjo, a primitive Bantu people
living on the slopes of Ruwenzori. Most of the cultivation was being
done by Batoro, while administration was largely in the hands of
Bahima, the tall Hamitics and former overlords. We found all of these
people most friendly and only too eager to dispose of ethnological
material, much of it superseded by changing customs.

loveridge: African itinerary and comments 197

Bugoye, Ruwenzori Mountain*. 0°17' N., 30°13' E. Alt. c. 5600 feet.

( lamped in the rest camp enclosure.

December 26 to 28th, 1938, and January 21 to 24th, 1939.

Occasional showers. We had left our thermometer hanging on a tree
in the heart of Kibale Forest so thereafter we were unable to record
temperatures.

After leaving Bundibugyo we drove over the northern end of the
Ruwenzori Range, which is about 30 miles in maximum width, through
Fort Portal, skirted the eastern flank of the 65-mile-long range, then
turned off the main road and took the side road to Bugoye where it
terminates.

Bugoye, at the southeastern foot of the range, is not only the type
locality of Chamaeleo h. ellioti, of which we obtained a good series, but
the jumping-off place for any ascent of the mountain (16,800 feet)
from this direction. This was the route followed by the British Mu-
seum Expedition of 1908, an expedition which resulted in several score
of new vertebrates. It was partly in the hope of securing topotypes,
at least of the reptiles, that we now planned to visit the two most
readily accessible sites occupied by Woosnam and his party.

We stayed at Bugoye only as long as it was necessary to get to-
gether sufficient porters for the ascent, while on the return journey I
was fully occupied with packing the specimens obtained on the moun-
tain before the lorry should arrive to pick us up.

Consequently little collecting was done in this uninteresting spot
whose surrounding slopes, eroded by constant rain, are clothed only in
sparse grass with thickets and, lower down, acacia or orchard forest.

Mubuku (Mobuku) Valley, Ruwenzori. 0°24' N., 30°0' E. Alt. 6800 feet.

Camped in deep forest between the Mubuku and Mahoma Rivers,
near foot of Nyinabitaba Ridge.

Arrived about noon on December 29, 1938, and left January 9, 1939.

Rain fell on every day except one. The worst aspect of the situation
was the absence of sunshine due to overhanging clouds which cast an
evening-like gloom and stillness over the forest for hours on end. Sun-
day was the best day with 7 hours of sunshine, on Monday about half
that amount, the rest averaged between 1 and 2 hours only. The tem-
perature at midday was probably from 55° to 66°. Under such condi-
tions collecting was difficult, birds were silent except during the brief
hours of sunshine when they were engaged in feeding on the forest
canopy well out of range. Drying of skins and other specimens became
a constant nightmare.

Despite the sodden state of the undergrowth only a toad and 3

198 bulletin: museum of comparative zoology

species of frogs were encountered, all the latter (Rana f. angolensis,
Phrynobatrachus graueri, and Hyperolius ? alticola) previously recorded
from the mountain.

Reptiles were limited to 5 species of which two were new for Ruwen-
zori, these were Cnemaspis a. elgonensis and Lygosoma g. graueri. This
little pentadactyle skink, together with the four-toed L. mcleagris, of
which we secured topotypes, were relatively plentiful. Topotypes were
also obtained of the chameleons (C. j. johnstoni and C. xenorhinus) ,
the latter a great rarity apparently dwelling in the forest canopy.
Naturally C. b. rudis, obtained by the British Museum Expedition at
10,000 feet, was not met with at the lower level but I was surprised to
see no Lacerta jacksoni, a species that has been taken at 8500 feet.

A pair of crimson-winged plantain-eaters, presumably Ruwenzoror-
nis, were seen near camp one day but out of gunshot in the tallest
trees. A mountain buzzard (Buteo oreophilus) visited our lonely camp
and fell to my gun, while topotypes of half-a-dozen passerines were also
collected.

Mammals were decidedly scarce, not a bat was seen, and, though a
net was spread across a suitable clearing, nothing was taken. Colobus
were seen only on the march to this camp and all the monkeys seen
were two solitary males of Cercopithecus m. stuhlmanni, the one col-
lected would be almost a topotype of the synonym carruthersi which
was first obtained a thousand feet higher up the mountain. During
our entire stay squirrels (Heliosciurus and Tcnniscus) were observed
four times only . Of three species of rodents preserved one was a topo-
type of the interesting mouse (Hylomyscus d. denniae). A red duiker
was seen once and heard a couple of times but attempts to collect it
failed. Not a trace of any carnivore was noted.

Mihunga Ridge, Ruwenzori. 0°21' N., 30°3' E. Alt. 6000 feet.

On the outward journey we had camped for the night beneath the
fig tree — on the upper Mihunga Ridge and collected a few specimens ;
returning we passed it to pitch our tents on the lower Mihunga Ridge
on the very site formerly occupied by the British Museum Expedition
in 1908.

Night of December 28, 1938. Then from January 9 to 19th, 1939.

We had fled from the Mubuku Valley camp on account of adverse
climatic conditions, now, though only a few miles distant as the crow
flies we seemed to be in another world. On this largely treeless spur,
bounded on the north by Weria Ravine, on the south by the Kanyon-
gorogoro Ravine, we were able to profit by the hot sunshine which
lasted for the first three days until 3 p.m. and was then followed by

loveridge: African itinerary and comments 199

showers. No rain at all occurred on the four days following while dur-
ing the last week, though the sky was often covered by fleecy clouds
through which the sun had difficulty in breaking, dark clouds formed
only in the late afternoon and then frequently dispersed without preci-
pitation in our vicinity, though at times rain might be seen falling
elsewhere.

Collecting was not confined to the 6000-foot ridge for excursions
were made to the forested heights a 1000 feet higher as well as to the
foot of the ridge where it tapered out into the swamps of the Mubuku
Valley. For descriptions of this and the last camp the reader is re-
ferred to Woosnam's account (1910, Trans. Zool. Soc. London, 19, pp.
5-24), and to some important comments by Allen and Loveridge
(1942, Bull. Mus. Comp. Zool., 89, p. 148).

Almost all of the amphibians (Bufo r. regularis, Leptopclis n. christyi
and Phrynobatrachus graueri) and most of the 9 species of reptiles
taken, came from the swamp about 500 feet lower than our camp.
Here, high in the papyrus, %were coiled the green vipers (Atheris n.
nitschei) topotypes of A. woosnami of which we had come in search.
Here also we obtained the first Uganda example of the dwarf chameleon
(Brookesia s. boulengcri).

Of 30 kinds of birds collected, 4 were topotypic, while each morning
our traps yielded some topotypic rodent until we had 10 of the species
collected at Mihunga by Woosnam and his associates; 3 others were
almost topotypic the types having been taken at higher altitudes.
This region appears to be as rich in mammals as it is poor in amphi-
bians for, in addition to the score of species collected, we found three
sleeping platforms built by chimpanzees in a tree in Weria Ravine.
The Bakonjo informed me that these platforms are made by itinerant
chimpanzees which come from the forests of the Mubuku in search
of bananas and, when benighted, construct a platform. We saw one
such individual while wTe were engaged in collecting in the swamp.

Nyakabande, Kigezi. 1°44' N., 29°45' E. Alt. 6925 feet.

Stayed at the commodious rest camp.

Arrived at noon on January 25 and left on 30th. Returning from
Mushongero on February 4, we had to wait for a lorry until the 8th.

Though there was relatively little sunshine, the weather was
generally fine except for a few heavy showers.

Nyakabande is situated in a lava-strewn plain, much of which is
industriously cultivated by the Banyaruanda who gather up the larger
blocks of lava and pile them on the periphery of each small plot, where
they serve as a windbreak. There is no water in the immediate

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vicinity, and perhaps it is just as well not to visit the foul pool from
which your water supply is likely to come. Presumably it was from
this pool that the four species of amphibia obtained here came.

Nyakabande is about six miles from Kisolo (the rendering given by
the Uganda Survey on its map A 530 of 1928, often misspelled Kisoro
or Kissolo), type locality of the toad I named Bufo r. kisoloensis,
whose validity can no longer be maintained as a result of the fresh
material obtained at Nyakabande and Mushongero.

The reptile fauna of these two localities is essentially similar as we
got only three species at the former not taken also at the latter, which
is reached by a very arduous climb over the mountains.

In the rest house grounds we found a weaver (Ploceus n. graueri)
nesting in a vociferous colony of the superficially similar P. c.feminina.
At Mushongero we obtained a topotype wagtail. (Motadlla c. wellsi)
and a pair of flycatchers (Alseonax a. ruandae) which race was de-
scribed from Bufundi on nearby Lake Bunyonyi.

At both localities the Banyaruanda apparently eat the huge mole
rats (Tachyoryctes ruandae), for they brought in as many as I would
take. The species was first collected on Mt. Muhavura which is in full
view of the rest house at Nyakabande. More valuable was an otter
(Lutra m. tenuis) which I shot in the lake near Mushongero, for it was
from this lake that Hinton described the synonym L. m. mutandae.

The main object of our stopover at Nyakabande, however, was to
purchase ethnological material from the teeming tribes inhabiting this
upland plain. Collectively known as Banyaruanda, they apparently
consist of a small admixture of Hamitic overlords, the cattle-owning
Batusi and Bahororo. The bulk of the population consisting of Bantu
agriculturists known as Bahutu, while a few semi-pigmy Batwa are
present, chiefly in the vicinity of the lakes.

Mushongero, Lake Mutanda, Kigezi. 1°46' N., 29°41' E. Alt. 5925 feet.

Stayed at the rest camp, which is situated on a little peninsula pro-
jecting from the east bank near its northern end.

Arrived late in the afternoon of January 30, and left early on
February 4th.

Frequent rainstorms swept across the lake whose mountain-girt
northern half appeared to attract and hold the heavy black clouds
which overhung it during much of our brief visit. When the sun was
hidden it was decidedly chilly.

Except for its extensive papyrus swamps, Lake Mutanda offers little
indication of its proximity to the equator. In fact with its numerous
tree-covered islets and purplish mountains its general appearance is

loveridge: African itinerary and comments 201

not unlike that of a Scottish loch. The brambles, bracken, and short,
wind-swept grass clothing the lower slopes of the mountains all suggest
that it was with good reason that these uplands have been called the
Switzerland of Central Africa.

As for the name Mushongero (misspelt Mushungero on all my
labels), I suspect that there is some connection between it and Mus-
hungwe, the Lugezi name for leach, for these loathsome creatures
swarm in the shallow waters in the vicinity of the rest camp. Crabs
(Ngara: Lugezi) were also not uncommon, and the series secured have
been described by my colleague, Dr. F. A. Chace Jr., under the name
of Potamon (Geothelphnsa) mutandensis.

The waters of the lake were teeming with Xenopus 1. bunyoniensis,
originally described from nearby Lake Bunyonyi, but search of the
papyrus both by day and night residted only in the capture of a single
example of the genus Hyperolius, and that unidentifiable! The most
perplexing aspect of the amphibian fauna was the intermediate condi-
tion of many Rana fuseigula, some, as one would expect from the ter-
rain, were typical, others showed the longer hind limb of the sylvicoline
race chapini. Three other ranids taken here also seemed to suggest
that some deforestation had taken place.

This was again the case with a couple of the dozen species of rep-
tiles collected. \Ye had gone to Mushongero primarily in search of a
blind snake obtained there by Col. Pitman; hoping that an adequate
series of the creature might settle its uncertain status. In this we were
successful, the 9 examples undoubtedly referable to Typhlops blanfordii
lestradci, originally described as a fidl species from nearby Ruhengeri
but obviously related to blanfordii of the Ethiopian highlands; another
link with the latter region is the presence at Mushongero of the little
slug-eater (Duberria I. abyssinica).

BELGIAN RUANDA

Kiraga near Kisenyi, Lake Kivu. 2°38' S., 29°1S' E. Alt. c. 5800 feet.

Camp was pitched in a plantation of eucalyptus through which the
road passes about three miles above Kisenyi, a township situated on
the shores of Lake Kivu at 4800 feet.

Arrived in afternoon of February 8, and left early on the 13th.

Some showers.

Most of our collecting at Kiraga was done along the banks of, and
in the ravine cut by, the Kisenyi River, which cascaded over falls just

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below our camp. The ravine was more or less choked by luxuriant
growths of grass and sedge, but in patches cleared for gardens we
turned over piles of vegetable debris and sought our quarry in the ex-
tensive plantations of bananas.

Only 4 species of amphibia and 6 of reptiles were taken, none of
especial interest with the possible exception of Lacerta jacksoni which,
in the absence of trees, has adapted itself to a terrestrial life. By far
the most important of half-a-dozen mammalian species were a pair of
skunk-like zorillas (Poecilogale a. doggetti).

As we were debarred from killing anything but "vermin" on ac-
count of our permit for scientific collecting not having arrived from
Stanleyville, we caught the first boat from Goma (Ngoma), type
locality for a burrowing viper (Atractaspis schoutedeni), which appears
to be based on a slightly aberrant example of the widespread A. irre-
gularis, which we got.

BELGIAN CONGO

Mamm Bay, Idjivi Island. 2°12' S., 29°0' E. Alt. 4788 feet.

Camped on the lawn of Mons. van der Berck v. Heemstede's estate.

Arrived at dusk on February 14 and left on 16th, returning March
6th.

Our arrival was greeted by torrents of rain and waves lashed by a
gale.

Wading in thigh boots among the sedges of the Bay, and aided by a
flashlight, I was able to capture topotypes of Ilyperolius kivuensis,
Icwidjwicnsis, kandti, and macrodactylus, and show that actually only
two sexually dichromatic species are present. The types had been
collected by the poet Kandt, who had made his home on Idjwi Island.

Upper Mulinga River, Idjwi Island, c. 2°8' S., 29°3' E. Alt ^500 feet.

Camped beside a footpath where it crosses a stream known as the
Upper Mulinga. This, I imagine, was about 900 to 1000 feet below
the 800 metre summit of the mountain, which dominates the island.

Arrived on February 16 and left at noon on March 6th.

The weather was very varied, we enjoyed much sunshine when it
would be quite hot; violent thunderstorms accompanied by lashing
rain were not uncommon, however, and always resulted in a consider-
able drop in temperature.

1 Not 4500 feet as printed on p. 149 of the report of Mammals, 1942, Bull. Mns. Comp. Zool.,
89, pj>. 147, 214. I am indebted to Dr. J. P. Chapin for pointing out this error.

loveridge: African itinerary and comments 203

Day after day I made excursions up the mountain to the extensive
remnants of magnificent forest, even then being destroyed by natives
contrary to regulations; policing of such remote spots being difficult.
Apart from some small patches of forest, our immediate vicinity con-
sisted of pasture land, millet fields, dense patches of sedge, and
swampy areas through which meandered small rivulets.

It was in this latter habitat that I collected most of the 8 species
of amphibia taken, but the choicest of all, a tiny, long-fingered male
Arthrolcptis xenochirus, was brought to me by a native lad. Only 2 of
the species were savanna forms.

Of the 25 species of reptiles collected, all but a fourth were of forest
association and included such rarities as Miodon g. graueri and Algi-
roides vaucreselli. The island, variously spelt Idschwi, Kwidjwi, and
Kidjwi by the Germans, is type locality for Mabuya m. kicidjwicnsis
and Lygosoma blochmanni. Over fifty of each were preserved, suffi-
cient to demonstrate that the former does not differ structurally from
true M. m. maculilabris, and that the latter is constantly three-toed.
The local chameleon appeared to differ sufficiently from its continental
congeners to be named C. d. idjwiensis.

Also new were 8 examples of a warbler (Apalis eidos), and we secured
topotypes of Barbatula kandti = Pogoniulus b. jacksoni, Colivs s.
kiwuensis and Spinus c. frontalis; in all 42 species of birds were col-
lected at this camp.

Two new races of rodents (Thamnomys v. kimiensis and Leggada b.
ablutus) were discovered, the former just behind my tent, the latter
differing from the typical race occurring on the Ruwenzori Moun-
tains. An interesting aspect of the mammalian fauna of this mountain
on Idjwi was that a third of the 21 species collected were referable to
races originally described from Ruwenzori; two others (Cercopithecus
m. schoutedeni and Lophuromys a. laticeps) were topotypes of local
forms.

TANGANYIKA TERRITORY

Ujiji, Kigoma District. 4°55' S., 29°42' E. Alt. 2800 feet.

Camped beneath the giant mangoes on the western fringe of the
town.

Arrived at noon on March 9, and left early on March 10th.

Occasional heavy showers following periods of sultry weather and
overcast skies.

As, from May 22 to 29, 1930, I had already visited this famous old

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Arab settlement on the shores of Lake Tanganyika, a description of
its features will be found in 1933, Bull. Mus. Comp. Zool., 75, p. 23.

The purpose of my visit was the same in both instances, viz. to ob-
tain examples of the rare Amphisbaena phylofiniens, known only from
the two cotypes described in 1905. On the first occasion I was unsuc-
cessful but learned from the natives that it was to be found in Ruanda,
a region of rice swamps bordering the Luiche River. Almost daily,
therefore, accompanied by two assistants, I tramped over to Ruanda,
pausing en route to inform each passer-by of our object. Not an am-
phisbaenid did we find, but as a result of our ceaseless talk 4 speci-
mens were brought in by natives. Since collecting all 7 East African
members of the family, I have come to the conclusion that the habitat
which these wormlike creatures require is one of moist sand or laterite
soil. There is no evidence which would justify the view that they are
survivals of a former forest fauna in this region.

While searching for amphisbaenids we secured a good series of a
Congolese skink (Scelotes t. hemptinnei) which had not previously been
taken in Tanganyika. Another first record for the Territory was a
"two-headed snake" {Chilorhinophis gerardi) though I had postulated
its occurrence at the southeast end of the lake in 1933. The Angola
race ornatum of Lycophidion capense was a further addition to the fauna
and, from a distributional point of view, an Aparallactus c. capensis is
interesting.

Additions to the fauna among the 9 species of amphibia taken, were
Rana m. venusta and Hypcrolius kivuensis; topotypes of H. udjijiensis
and argentovittis were also captured.

Kitaya, Southern Province. 10°40' S., 40°11' E. Alt. 300 feet.

Tents were pitched on the rest camp site on the bank of the Rovuma.

Arrived at noon on March 24th and left early on April 7th.

Torrential rains fell during our stay, necessitating the employment
of porters for the first nine miles of the return journey to Mikindani,
i.e. until past a depression of water-logged black cotton soil.

Kitaya is a village, and Liwale headquarters, on the north bank of
the Rovuma, (Rowuma; Ruvuma) River, fifteen miles inland from
Rovuma Bay. The River forming the southeastern boundary between
Tanganyika Territory and Mozambique. Livingstone safaried there
from Mikindani in 1866; the two places are now connected by a very
rough motor track of about 36 miles. The vegetation is typical of the
coastal belt; baobabs, orchard forest, and rank grass on the higher
ground, which is very sandy; rice fields and sedge (Setaria palmifolia)
fringed swamps on the lower.

loveridge: African itinerary and comments 205

Huge leeches are present in the swamps, ticks in the long grass.
Aedes, Glossina, Haematopota, Stomoxys, Tabanus, and many other
biting flies were an ever-present nuisance.

The purpose of our visit was to secure topotypical material of three
species of sedge frogs {Hyperolius eitrinus, microps, and Megalixalus
fiavomaculahis) collected by Kirk when he accompanied Livingstone.
In this we were eminently successful and able to prove that the last
named is really a Hyperolius, having a horizontal pupil in life. In
addition a series of an undescribed race of this genus (//. p. rovumae)
were collected and described.

We had to depend Aery largely on our own efforts, for the local
people — Konde and Yao — were strangely disinterested in bringing in
reptiles or mammals, in fact supplied only two of the latter. Crocodiles
as man-eaters play a prominent role in local village life. Soft-shelled
turtles (Cycloderma frenulum) were laying and in consequence may
have been more conspicuous than would normally be the case. The 13
species of snakes might almost have been selected as representative of
the most widespread African forms! Except for Ichnotropis squamuhsa
the dozen species of lizards were likewise of little interest.

Birdlife was wonderful, being particularly rich in non-passerine
species such as parrots, hornbills, wood hoopoes, cuckoos, and wood-
peckers. In all 61 species of birds and a dozen different kinds of mam-
mals were collected during the 13 days spent here. Of the latter a
molossid bat (Mops angolensis orientis) was described as new.

Mikindani, Southern Province. 10°17' S., 40°7' E. Alt. 20 feet.

Tents were pitched about two miles north of the centre of the town-
ship, which gives its name to Mikindani Bay on the southeast coast of
Tanganvika Territorv, on a little rise to the left of the mainroad to
Lindi after one passes the Government pumping station. Another
noisy pumping station engine has given the name of Mchuchu to
this area.

First landed at Mikindani on the night of March 22 and accom-
plished some collecting on the following evening before leaving for
Kitaya. Returning from the latter place on April 7 in torrential rain,
we just managed to get the tents pitched before darkness fell on the
most cheerless conditions of the whole trip. Left by dhow for Lindi,
the road being under water for miles, on April 24th.

Fully 0 inches of rain fell during our sixteen days stay, on some days
continuing from dawn till dusk. The greatest single precipitation
recorded was 1 3/16th inches on April 10, for this information and
other kindnesses I am indebted to Mr. E. A. Leakey, District Officer.
The average annual rainfall is 36 inches.

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Such conditions naturally retarded outdoor studies to some extent.
Most of our collecting was carried out in the eastern environs of the
township and relatively little in the immediate vicinity of camp.
Though the low ground surrounding the latter was largely under water,
the rainy season was already so well advanced that spawning was over
for the majority of frogs, which were already widely dispersed. Of
the 13 species collected only one (Arthrolcptis xenodactylus) has any
claim to forest associations.

Such primeval forest as may have been here long since disappeared,
but the numerous mango and other trees, to say nothing of baobab and
coconut, result in* providing conditions acceptable to some sylvicoline
species. Of the 31 species of reptiles taken, however, only a cobra
(Naja melanoleuca), is a western forest form, the 7' 6" specimen pro-
viding the most southeasterly record for the species. Topotypes were
collected of the only two lizards (Lygodactylus g. grotci and Amphis-
baena orientalis) described from Mikindani.

Of the 43 species of birds collected, topotypes were shot of Francoli-
nus h. grotci, Lagnosticta r. rcichcnowi = haematocephala, and Uraegin-
thus b. mikindaniensis = niassensis, while of 3 other races described
from here we had obtained specimens from nearby Kitaya and Nch-
ingidi.

Only 7 kinds of mammals were obtained, all typical of the coastal
zone, the bat Triaenops afer being, perhaps, the most interesting.

Mbanja, Southern Province. 9°24' S., 39°45' E. Alt. Sea level to 400
feet.

Camp was made at the edge of Mitonga (Metonge) landing field
('aerodrome'), circa 375 feet.

Arrived at noon, by truck from Lindi, on April 25, and left at 8 a.m.
on May 6, 1939.

Heavy showers occurred during the first week; the second was prac-
tically rainless, the heat tempered to some extent by the southeast
monsoon, until we were struck by a gale of wind and rain on our last
night.

Mbanja (Mbanya), about 10 miles north of Lindi, is a small village
situated on a tidal estuary and almost surrounded by mangrove
swamps. The chief and his people were most friendly and helpful.
While the village itself is on clays and coral rock, the valley at whose
mouth it lies is largely composed of black cotton soil and rich mud ex-
tensively cultivated by the industrious inhabitants. Their principal
products being ground nuts, potatoes, and mahoga, with a few paupau
trees and coconut palms about each hut.

loveridge: African itinerary and comments 207

My camp, on higher ground, was largely surrounded by orchard
forest heavily interspersed with mango trees and waist-high grass
whose barbed black seeds cause considerable discomfort by working
through clothing and even puttees. To the west lay dense scrub in-
habited by very wary squirrels and blue monkey, for the cry of the
latter was heard towards sunset on several occasions. To the east,
i.e. between camp and the coast, were native gardens and the exten-
sive Kikwetu Sisal Estate. All this country is composed of bright red
and very porous soil derived from eroded limestone.

It is in this soil that Amphisbaena ewerbecki, described from here by
Werner in 1910, occurs. Our object in coming to Mbanja was solely
to get a series and in this we were entirely successful. The soil which
furnished a congenial habitat for the amphisbaenid was favoured by
six species of fossorial snakes which we collected in addition to 9 other
kinds of ophidia, a dozen lizard forms, and 36 of birds.

Game was allegedly shot out by the Germans, who maintained a
big camp near here during 1914-1917, but wild pigs were common. At
night a solitary little antelope emerged from the scrub to feed on the
foliage of the ground nuts. Other animals seen or heard, but not col-
lected, were red elephant shrews, galagos, baboons, hares, ratel and
jackal. Man-eating lions had been causing numerous deaths quite
recently in neighbouring villages.

Rondo Forest, Southern Province. 10°8' S., 39°12' E. Alt. 2700 feet.

Camped at Nchingidi, the name given to a clearing at the forest
edge approximately three miles from the nearest scattered native huts.

Arrived just before noon on May 9, and remained until the 21st.

Each evening at varying times from sunset (6 p.m.) till 11 p.m., a
succession of mist clouds blew in from the ocean and up the face of the
escarpment (at whose edge my tent was pitched) to condense on grass
and trees. The latter literally 'rained' upon my tent with every gust
of wind. At daybreak these mist clouds hemmed us in, completely
shutting out the view of the opposite escarpment, invariably persist-
ing till 8 a.m., more usually 9 a.m., occasionally even until noon. In
addition about 4 or 5 p.m. there were sharp showers, while sudden and
unexpected storms of rain swept the plateau at uncertain intervals,
and frequently heavy downpours occurred at night. The nights were
always cool but when the sun did break through by day it was very hot.

Unfortunately an error deprived me of my quinine, and repeated
drenchings when far from camp brought on a fever which, so far as I
was concerned, halted collecting during the last week.

This open forest is situated on the waterless Rondo Plateau, about

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25 miles sothwest of Lake Rutamba, (Lutamba), itself some 25 miles
by road slightly southwest of Lindi. The curious thing about this
forest, of which mvidi is the most important constituent, is the entire
absence of standing water. Even the heaviest downpours immediately
disappeared in the porous sandy soil. To obtain water for domestic
purposes, the local natives (Mwera) made a daily journey of three
hours (there and back) to the foot of the escarpment. It is not alto-
gether surprising that with water so scarce these people should con-
sider washing superfluous. Though very friendly, these Wamwera
only began to busy themselves with bringing in specimens a day or
two before our departure.

Despite the absence of water, 5 species of amphibia, all previously
taken on the Uluguru Mountains nearly 300 miles to the north, were
collected. Of these a tiny toad (Bufo micranotis rondoensis) differed
sufficiently to warrant description.

This isolated plateau had produced 3 species of fossorial reptiles
(Amphisbaena rondoensis, Melanoseps a. rondoensis, Typhlops t. ron-
doensis) which it was necessary to describe as new. Here also we
found Chlorophis maerops, Bitis gabonica, and Brookesia brevicaudata
which occur on the Usambara Mountains 350 miles to the north ; even
more surprising was an undoubted Aparallactus jacksonii of Kiliman-
jaro. The remaining reptiles — 12 kinds of snakes and 10 of lizards
were representative of the coastal plain herpetofauna.

In the past, principally during the Great War, the forest suffered
heavily from. native incursions. Subsequently these refugees were re-
moved but evidence of their destructive occupation were to be seen
in the numerous clearings, some so eroded as to be semiarid areas with
only the scantiest covering of grass, others grass- or bush-grown and
inhabited by such non-sylvicoline, house-dwelling lizards as Hemi-
dactylus mabouia and Mabiiya striata, which I assume to have been
imported by human agency.

Bird life, though not abundant, presumably owing to the absence
of standing water, held promise of being interesting so that it was a
disappointment to go down with fever just as I was turning my atten-
tion to the avifauna. A beautiful roller {Eurystomus glaucurus) of
Madagascar was preserved, while the records of the green -headed
Oriole (0. chlorocephaius) from Mt. Chiradzulu, Nyasaland and the
Usambara Mountains are at last bridged by its occurrence on the
Rondo Plateau. Two rare flycatchers (Batis reichenoioi and Erythro-
cercus I. thomsoni), originally described from Mikindani and the
Rovuma River respectively, were also among the 25 species obtained
at Nchingidi.

loveridge: African itinerary and comments 209

Doubtless the water supply is also the reason for the scarcity of
mammals, of which only 9 species were taken, the most interesting
being Rhynchocyon p. melanurus, originally described from Lindi. I
heard, but never saw, blue monkey, bushbuck, and a small duiker
which the natives called naunde.

Lindi, Southern Province. 10°0' S., 39°14' E. Alt. 50 feet.

In Lindi Hospital (one week) and at the Beach Hotel (one week).

Arrived on May 22 and sailed on June 5th.

The rains, which average 34 inches per annum, were over, and the
clay soil already baked hard by a tropical sun.

Lindi, headquarters for the Province of the same name, is situated
in Lindi Bay between Kilwa and Mikindani. A population of nearly
4000, and the excellent hygenic conditions of the township, rendered
collecting difficult during the week that I daily awaited the arrival of
a steamer. All the same we preserved 15 species of amphibia and
reptiles, 16 of birds, and 5 of mammals.

On our very first hunt, however, we uncovered the oriental blind
snake (Typhlops braminus), the fourth and fifth examples to be taken
in the Territory. T. s. mucruso was the only other snake obtained
though bundles of thatching grass, piles of palm fronds, and heaps of
rubbish, were turned over a wide area. Amphisbaena cwcrbccki was
the only lizard of interest. An intelligent fisherman told me that four
turtles are occasionally encountered in the Bay, he described the luth,
loggerhead, hawksbill, and green turtle. ,

Siga Caves, Tonga Province. 5°6' S., 29°4' E. Alt. c. 150 feet.

Camp was pitched on the trail leading to the caves but about a mile
from the entrance.

Arrived on June 7 and remained until the 17th.

A few heavy rainstorms occurred at infrequent intervals, the rainy
season being over.

The Siga Caves, I took the name from the government signpost
erected at the turn off from the main Tanga to Mombasa road, have
nothing to do with the Sigi River a few miles to the north. They are
sometimes called the Amboni or Mkulumusi Caves for the main en-
trances are less than fifty yards from the crocodile-infested Mkulumusi
River. The numerous caves are waterworn and in past times un-
doubtedly formed an underground channel for the river. There are
dubious stories of the caves extending for a mile, but at the time of our
visit every passage was flooded to within a hundred yards of the
entrance.

The fifty acres of forest which surrounds them has suffered con-

210 bulletin: museum of comparative zoology

siderably, in fact it seemed to me that little remains but a scattering
of fine trees surrounded by secondary growth and much scrub. I
might add that the caves are held in superstitious veneration by the
natives, and propitiatory rites, of which I have given a brief account
(1940, Scientific Monthly, 51, pp. 22-35) were held there at the time
of our visit. I imagined that the presence of this "spirit" was the
reason for the absence of native squatters : later I learned that during,
and immediately following, the Great War, natives had moved in but
were turned out again by the Administration on account of their reck-
less destruction of trees.

All 11 species of amphibia, as well as 13 kinds of reptiles obtained at
Siga, were typically coastal plain. One gecko (Cncmaspis a. africanus)
was definitely sylvicoline, while five others proved of assistance in de-
fining a new coastal race named Hcmidactylus t. barbouri.

No serious attempt was made to collect birds in so well-worked a
region, and only 8 species were shot, the choicest being a pair of
migrant Malagasy egrets (Egretta g. dimorpha) and my first bat-eating
hawk (Machaerhamphus a. anderssoni).

The latter was, of course, attracted by the thousands of bats which
made the caves their home. Of these bats, 96, representing 4 species,
were collected ; they ranged in size from the huge Hipposideros g. gigas
to the modest Miniopterus minor. Galagos (G. c. lasiotis) were also
plentiful, their strange cries, mingling with the staccato bark of hyrax
and the hooting or screeching of owls, rendered our nights the noisiest
of the entire safari. The 10 species of mammals, all troglodyte or
arboreal except for a spiny mouse {Acomys w. wilsoni), were of coastal
rather than of forest affinities.

Amboni Estate, Tanga Province. 5°3' S., 39°3' E. Alt. 300 feet.

Our tents were pitched beside a small area of forest, largely second-
ary, which is being carefully preserved by the Estate Management.
As Amboni Estate covers nearly 80 square miles, the precise location
may be somewhat important, we were located about a mile above
Mabokweni Village and surrounded on three sides by sisal planta-
tions.

Arrived on June 17 and remained until the 27th.

Rain occurred in scattered showers on about four cloudy days.

It seemed strange that frogs should be assembling at the conclusion
of the rainy season, yet a visit to a rice swamp close to Mabokweni
Village on the evening of our arrival, resulted in the capture of 154
polypedatid frogs of 9 different species, the females of Hylambates
maculatus certainly were gravid and about to spawn.

loveridge: African itinerary and comments 211

Between this swamp and our camp was an extensive area that had
been under sisal since 1913. It had recently been cleared and tractors
were now engaged in spreading the vegetable debris which had been
piled in long rows and left to rot for two months. By following the
twelve-ton tractors to and fro for three days, two species of ranid
frogs, 5 of lizards, and 11 of snakes were secured. The latter were
largely of fossorial types, among them a burrowing viper (Atractaspis
bibronii) which conformed to the description of katangae of the south-
ern Belgian Congo! Only one frog (Rana o. gribinguiensis) and one
snake (Calamclaps u. warreni) could be cited as largely sylvicoline.

Nor did the forest appear to have any significance as a refuge for
surviving forest forms of reptiles, the 5 species taken within its con-
fines being coastal bush or savanna species. Hornbills, both Bycanisies
and Lophoceros, were much in evidence, but no small birds were shot
though several hours were spent in looking for them.

Jumping shrews (Petrodromus s. sultani) were not uncommon on the
outskirts, and galagos of two species (G. c. lasiotis and G. s. zanzibari-
cas) were present, the latter extremely plentiful. The place was a
refuge also for a band of colobus (C. p. palliatus) and some blue
monkey (Cercopithecus m. monoidcs) though there was no thought of
protecting the latter for whose heads a reward was offered as they
attacked the sisal shoots. Bats, a single red squirrel, and a bushbuck
complete the list of mammals seen in the forest, though other species
were collected in the surrounding plantation.

Magrotto Mountain, Tonga Province. 5°8' S., 38°45' E. Alt. 2500 feet-
Camped on a low hill half-a-mile as the crow flies west of the factory
of MagrOtto Estate.

Arrived on June 27 and remained until July 20th.
The first five days were largely overclouded, raw, damp, and chilly.
Cloud-mist like a dense fog, swept into the valley each evening about
4 p.m. remaining until 9 or 10 the following morning when dispersed
by the frequent rainstorms which swept across the hills. With the
advent of the new moon the vapour was reduced to capping the higher
forested ridges ; the hours of sunshine increased and for an entire week
there were fewer showers. During the last week, however, the weather
remained dull with frequent rainstorms. Average annual rainfall 75
inches !

Magrotto Estate, which formerly occupied about 5000 acres of hill-
tops capping the mountain, is reduced to half that size today. The
more than a million coffee trees of German times have given place
to the West African oil-palm, the only plantation of this palm in all

212 bulletin: museum of comparative zoology

East Africa. My camp among these palms was almost surrounded by
a horseshoe-shaped ridge that was largely forested, a gap towards the
south admitting the clouds of vapour which, during the southwest
monsoon, are an almost regular evening phenomenon. Just below the
western foot of the camp flows a river which rises nearby. To the
north is a swampy bottom, part of whose smothering mat of vegetation
we cleared in our search for frogs.

Prior to 1900, or thereabouts, the whole mountain was heavily
forested until much was cleared for coffee planting. During the Great
War natives moved in on the crown land and destroyed almost every
tree. The hillside directly opposite my camp had been cleared and
rows of Grevillea planted as shade trees for the coffee. When the Estate
was abandoned during the war, native trees, protected by the Grevillea,
sprang up and are now between 60 and 70 feet in height. I mention
this as representing the most amazing come-back of forest which I
have seen anywhere in East Africa. The forest-floor conditions ap-
peared indistinguishable to me from those in adjacent virgin forest.

Despite this, however, animal life, which in species closely resembles
the fauna of Amani at 3000 feet in the Usambara Mountains, and only
20 miles distant across the plain, was decidedly scarce; birds were con-
spicuously absent. In part, of course, this may be seasonal, for the
three weeks spent at Amani in 1926 were during the November rains
when amphibia were breeding, hence their predators more in evidence.
The three weeks spent on Magrotto were immediately following the
coldest months in the year when the temperature falls 57° F. (14° C.)
and a proportion of poikilothermous vertebrates may be assumed to
be quiescent.

Another factor, though only affecting the snake census, might be
found in the composition of the plantation personnel. At Amani the
'hands' were largelv of the Nvamwezi tribe, who are notoriouslv less
fearful of these reptiles. At Magrotto the labour was largely drawn
from local people, who, though helpful, and willing to bring in such
snakes as they came across, displayed no great enthusiasm.

At Magrotto 21 species of amphibia were collected of which only 3
had not been taken at Amani, 1 of these was the forest-edge form
(Rana m. venusta) of the widely distributed savanna species; signifi-
cantly enough both the others were recent invaders from the coastal
plain, i.e. Rana o. oxyrhynchus (instead of the forest-edge R. o. gribin-
guiensis taken at Amani) and Arthroleptis s. stenodactylus (instead of
the sylvicoline A. s. lonnbergi taken at Amani). Three remaining
species taken at Amani though not encountered on Magrotto, were

loveridge: African itinerary and comments 213

Xenopus and Hyperolius forms typical of the coastal plain and which
almost certainly will be found on Magrotto. A good series of topotypes
of H. substriatus (= puncticulatus) were preserved.

When we examine the composition of Magrotto's reptile fauna, of
which 20 kinds were collected, we encounter a similar situation. In-
stead of Typhlops p. gierrai, whose presence one might reasonably have
expected, we encountered only the typical form; as for Neusterophis
olivaceus, the montane uluguruensis occurred in almost equal propor-
tions to the typical savanna race. To sum up, for both snakes and
lizards, precisely 50% were sylvicoline, the rest savanna, reflecting
the fact that, though we spent more time hunting in the forest, reptile
life was more conspicuous in the plantation where immigrant forms
were alike supplanting the forest fauna in the zoological and botanical
realms.

As already stated, forest birds were scarce at the time of our visit,
only 6 or 7 being obtained, of which half were bulbuls. To these might
be added the great eagles (Stephanoaetus coronatus) seen. Of the species
collected 1 (or 2) were described from the nearby Usambara Moun-
tains, 4 from Kilimanjaro, and 1 from Mt. Elgon. A total of approxi-
mately 25% of the species being sylvicoline.

Similarly with the mammals not more than 25% of the 16 species
collected were forest forms in its restricted sense, though the propor-
tion might be raised almost to 50% by the inclusion of certain doubt-
ful creatures like Heliosciurus u. undulatus of Kilimanjaro which, like
the galagoes and monkeys, are arboreal and probably as much at
home in the coastal bush as in virgin rain forest.

Tanga, Tanga Province. 5°40' S., 39°7' E. Alt. 50 feet.

At Tanga Hotel prior to embarkation.

Arrived at noon on July 21 and left on the 23rd.

Fine and hot. Average annual rainfall 59.24 inches.

Spent my last afternoon in Tanganyika searching for Typhlops
platyrhynchus, of which Tanga is type locality, in the sandy area from
which the coconut palms had been removed in order to prepare the
site for a military landing field. All that we got were 3 species of frogs
and 2 of geckos, all typical of the coastal plain life zone.

KENYA COLONY

Likoni, Set/die Province. 4°5' S., 39°39' E. Alt. 50 feet.
On board R. M. S. Dunbar Castle.
Docked alongside Kilindini Wharf July 24 and sailed on the 26th.

214 bulletin: museum of comparative zoology

Showers and sunshine. Rainfall very variable, but the annual
average about 51 inches.

Likoni is a ferry landing on the mainland opposite Kilindini, Mom-
basa Island, the second locality in bold face type appearing on many
labels. It is a region of old coconut plantations and typical coastal
vegetation. I spent two mornings there in search of additional ma-
terial of a new gecko (Hemidactylus t. barbouri) which I had taken
previously at nearby Changamwe, and which I purposed describing.
In addition to this and other lizards we captured the first three
examples of a skink (Riopa pembanum) which I had ever collected, the
species being unknown from the continent until relatively recently.
A frog, two pigmy mice (Leggada b. vicina) of a race described from
Takaungu a few miles to the north, and a good haul of invertebrates
comprised the final spoils of a trip which had lasted nine months.

PLATES

PLATE 1

Loveridge — African Itinerary

PLATE 1

Map showing Principal Collecting Localities

1938

Landing at Mombasa (25.x), except for a stopover at Naivasha and Kinan-
gop (26-31. x), Loveridge proceeded by rail direct to Jinja (1-5. xi). Thence to
Mabira Forest (5-21.xi), Budongo Forest (22.xi-7.xii), Kibale Forest (8-19.xii),
Bundibugyo near Bwamba Forest (19-26. xii), Bugoye, foot of Ruwenzori
Mountains (26-28.xii) and Mubuku Valley at 7000 ft. (29.xii-).

1939

On leaving Mubuku (-9.i) Loveridge descended down the vallev to Mihunga,
circa 6000 ft. (9-21.i), then back to Bugoye (21-24.i), Nyakabande (25-30.i),
Mushongero (30.i-4.xi), returned to Nyakabande (4-8.U); Kisenyi (8-13. ii),
Goma (13-14.ii), Mamvu on Idjwi Island (14-16. ii), Upper Mulinga on Idjwi
(16.ii-6.iii), Uvira (7-8.iii), TJjiji (9-16. iii), Dar es Salaam (18-19.iii), Mikin-
dani (22-24.iii), Mbanja (25.iv-6.v), Lake Rutamba (6-8.v), Nchingidi
(9-21.v), Lindi (22.v-4.vi), Siga Caves (7-17.vi), Amboni Estate (17-27.vi),
Magrotto Mountain (27.vi-21.vii), Tanga (21-23.vii), Kilindini (24-26.vii).

BULL. MUS. COMP. ZOOL.

Loveridge. African Itinerary and Comments. Plate 1

UGANDA.-

'■•'BuJorrao

h> Forest i ^t
rJ*/CibaU F°rzZ~Apt&

Muill/iu Valley Bl Sf JMJ\

?' -Cb* Mabira '■

*■*«' MutaXijS—-^
Mil jCpn>V>°»S"°
Nyahabaniei -f(abale

Lake KlvJikk.%-

Costermansn-

Uvira

BELGIAN
CONGO

K EN YA
COLONY

TAN G A
TE R R

«0C

"aS"~vtto /t,f''WW«tJo MOMBASA
Ambon! EstaleY^f^t

Stga Cai/es'T Q

-v ( 9

ZIBAR
ES SALAAM

'Mindani

Scale in Miles

100

_l_l

200

PLATE 2

Loveridge— African Itinerary

PLATE 2

Fig. 1, Transport by water — Canoes on Lake Mutanda

These dugouts, waiting to take us from Mushongero to the south end of the
Lake, speak volumes for the patient toilers who, with adze or other simple
tool, gouged out the hard timber from the fallen tree. Moreover, as no trees
of sufficient height to provide sixteen or eighteen-foot canoes are to be found
within many miles of the Lake, these incredibly heavy craft had to be pushed
uphill and down dale on rollers by human muscle alone.

Fig. 2. Transport by Land — Our Lorry at Mubango

African travel today involves increasing use of modern methods of transport,
resort to head-porterage being reserved for roadless regions or where the ascent
of a mountain by native track is called for. In modernizing Africa motor-
driven vehicles will soon be claiming precedence over predators and snakes
as the major menace to life and limb. The photograph depicts our native-driven
Mercedes-Bentz, which at the time was carrying my six boys atop a load con-
sisting of a ton of camp and collecting equipment, irresolutely resting on
marshy ground where faulty driving had landed her at the very start of our
'safari!'

BULL. MUS COMP. ZOOL.

Loveridge. African Itinerary and Comments. Plate 2

PLATE 3

Loveridge — African Itinerary

PLATE 3

Fig. 1. Prospecting For a Camp Site — Forest-edge, Budongo

Budongo Forest, which covers about 180 square miles, gathers about its
fringes a dense and often impenetrable undergrowth of bush. Within, however,
where the unbroken forest canopy, two-hundred feet overhead, prevents
shrubs springing up among the mighty buttress roots, one can wander at will
over the thick layer of leaves which carpet the forest floor.

Fig. 2. Tractor and Tramway are Employed for Logging at Bisu

The absence of paths and water drove us unwillingly to camp at Bisu, in
close proximity to the Buchanan Saw Mills. There access to the forest was
assured by a track cut for the tramway depicted above. Unfortunately the
noisy tractor and its train, which made the three-mile run through the forest
every twelve hours, had driven elephant, buffalo, and other large mammals to
seek quiet elsewhere.

BULL. MUS. COMP. ZOOL.

Loveridge. African Itinerary and Comments. Plate 3

MHMETttBl^BUHB

PLATE 4

Loveridoe — African Itinerary

PLATE 4

Fig. 1. Native Path through Mabira Forest

Paths were few and far between, yet the undergrowth was so impenetrable
that they provided the only means for reaching the deeper forest to sample its
denizens. While monkeys and squirrels fell victims to this inquisitiveness,
parrots screeched or clambered about in the tree-tops, where, with a host of
smaller birds feeding in the forest canopy, they enjoyed complete immunity,
being out of range of gunshot.

Fig. 2. A Semi-pigmy of Baamba or Batwa Stock

The Baamba clans inhabiting the Ituri Forest region northwest of the
Ruwenzori Mountains, closely resemble their Batwa kinsmen of the Congo
forests. Attired only in a civet skin, the sturdy subject of the photograph
presents a marked contrast to the tall, cotton-clad Muganda on the forest
path in Mabira.

Fig. 3. Family Group at Bundibugyo in Toro

Yet another popular style in dress is exhibited by this Bantu family relaxing
after their early morning labours in the hot region just above the Semliki
Valley. Hundreds of natives representing many tribes, visited our camp at
Bundibugyo. If I am not mistaken those shown are Batoro, the agricultural
middle-class members of this society, superior to the hunting Baamba from
whom they obtain meat in exchange for cereals, yet themselves formerly sub-
ject to their pastoral overlords, the Hamitic Bahima.

BULL. MUS. COMP. ZOOL

Loveridge. African Itinerary and Comments. Plate 4

r-£.j

■&

■

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 6

NOTES ON LYCAENID BUTTERFLIES

By Harry K. Clench

MUS. COMP. ZOOL
LIBRARY

SEP 2 2 1964

HARVARD
UNIVERSITY.

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

July, 1944

PUBLICATIONS
OF THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
investigations by the Staff of the Museum or of reports by spec-
ialists upon the Museum collections or explorations.

Of the Bulletin, Vols. I to XCIII, and Vol. CXIV, No. 1, 2,
3, 4 and 5 have appeared and of the Memoirs, Vol. I to LVI.

After 1941 no more Memoirs are to be published.

SEP 2 2 1964

HARVARD
Bulletin of the Museum of Comparative Zoology rsiTY

AT HARVARD COLLEGE

Vol. XCIV, No. G

NOTES ON LYCAENID BUTTERFLIES

Bv Harry K. Clench

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

July, 1944

JUL |9 1944

L I B K A K *

No. G — Notes on Lycaenid Butterflies
By Harry K. Clench

a. THE GENUS CALLOPHRYS IN NORTH AMERICA

While working on a revision of the genus Indsalia Scudder sueh
frequent recourse was had to the various American species of Callo-
phrys that it was thought advisable to prepare a brief review of them.
Unfortunately, time was all too short to do the job properly, and
genitalic examinations were not made. These notes, however, were
gotten together, in the hope that they would facilitate further work
on the genus. Such future work should certainly include a genitalic
study of all forms involved.

The first and only real revision of the American Callophrys was that
of Barnes and Benjamin' , who placed the genus on very solid footing,
especially as compared with its previous state. It was through this
paper that our first accurate knowledge of the limits and variabilities
of the several species was obtained. There have remained, however,
several points that need clarification. Probably the most important
of these is the interrelationship of the various Californian forms. The
arrangement adopted by Barnes and Benjamin (dumetorum in the
north, with subsp. perple.va in the south) is too simple, and although
it at first appears logical, subsequent research has proved it false.
There are at least three forms, and very likely several more, in Cali-
fornia that have been going under the names dumetorum and perplexa.
What has been commonly passing as true dumetorum is something
else, while true dumetorum itself appears to be rather poorly known.
These will be discussed in greater detail under the species concerned.
Unfortunately, the problem in California has been only partly solved,
the material at hand indicating much further work to be done, but of
itself insufficient to do it.

In addition to this, there has been a new species (comstocJci) pro-
posed since the work of Barnes and Benjamin, and in the present
paper a second novelty, a new subspecies of affinis, is added.

Finally, there are a number of new and interesting records, exten-
sions of ranges, particularly of affinis and sheridanii.

The genus in North America appears to be confined exclusively to
the Rocky Mountains and the area westward to the Pacific. It ex-
tends from northern Mexico in the south to Alberta and British

U923, Contrib. Nat. Hist. Lep. North America 5, p. 61-67.

218 bulletin: museum of comparative zoology

Columbia1 in the north. The species appear to be generally mountain-
loving, although they are frequently (perplexa, for example) taken at
low elevations.

The paucity of species in Asia and Europe and their close inter-
similarity, as compared with the relatively large (6) number of species
and their diversity here in America, would suggest that the Palae-
arctic species were derived from American stock. The origin of this
American stock, however, is quite another question, and one foolish
to speculate upon, in view of our present ignorance of much of the
pertinent details.

The author wishes to thank the several individuals and institutions
who have greatly assisted in this review. The American Museum of
Natural History (AMNH) very kindly loaned some material for
study. The Carnegie Museum and the U. S. National Museum
checked the types of species in their collections. Mr. Robert G. Wind,
of Berkeley, California, was very kind in sending me for study a large
amount of exceedingly interesting material, including a number of the
complex viridis-\\ke specimens from the Californian Sierras, and other
northern Californian localities. Mr. Charles L. Remington also lent
some very interesting material which was of the utmost help in pre-
paring the paper. Mr. L. P. Grey gave the author all of his Callophrys,
which included several very interesting things. The collection of the
Museum of Comparative Zoology (M.C.Z.) has, as in times past,
always been available to me through the kindness of Mr. Banks. This
collection contains much material of interest and assistance. My own
collection has been drawn on wherever possible, being designated
(H.K.C.).

Genus Callophrys Billberg

1820, Enumeratio Insectorum, p. 80. Genotype, Papilio rubi Linn. (Palae-
arctic).

Key to species

1. Fore wing underside with green covering wing down to C112 or 2A 2

- This green restricted to a more or less broad costal and outer marginal
border, leaving a large internal area of gray or fulvous 6

'The two papers on the faunae of these provinees (K. Bowman, 1919, Annotated Cheek List
of the Macrolepidoptera of Alberta, Alberta Nat. Hist. Soc., Red Deer, Alta., 16 pp.; E. H.
Blackmore, 1927, Check-list of the Macrolepidoptera of British Columbia, C. F. Banfield,
Victoria, B. C 47 pp.) each list Callophrys dumelorum. Just, what they meant is not known.
Indeed, it coidd have been almost any species in the genus, but most logically affinis or shcri-
danii. Beferences to these papers, as well as to a number of others, were omitted due to this
and similar lack of precise information, so common in this genus.

clench: notes on lycaenid butterflies 219

2. Hind wing below immaculate, or at most with a faint indication of a white

line — usually a discal row of obscure white dashes 3

- This wing with either a prominent white line, or with a series of distinct,

pure white dashes or dots 4

3. Fulvous above in both sexes widespread, dominant; underside apple-

green a. ajfmis

- Fulvous above reduced, frequently (usually?) absent in the male; green

below purer, inclining even to bluishness affinis washingtonia

4. Underside of hind wing with a solid line of white 5

- This surface with a row of prominent white spots viridis

5. Line on underside of hind wing heavy, inwardly edged with a pronounced

band of black s. sheridanii

- This line thin; the black faint or absent sheridanii neoperplexa

6. Underside of hind wing nearly immaculate, or with a row of white spots

of varying number 7

- This area with a crooked white line 8

7. Outer margin of fore wing evenly convex apama homoperplexa

- This margin convexly angled at M2 or thereabouts, and straight, or

slightly concave thence to the inner angle 9

8. Line on underside of hind wing basally edged with black, then fulvous

a. apama

- This line with no fulvous comstocki

9. Costa of fore wing below edged with fulvous dumetorum perplexa

- Costa of this surface not edged with fulvous (or at most only very slightly)

d. dumetorum

Callophrys dumetorum Boisduval

This species ranges from northern Mexico (Baja California) north
beyond San Francisco. The northern limits of its range are still un-
defined. There are apparently two subspecies of this, one very widely
known (perplexa), inhabiting the lower part of its range, and the other
(typical dumetorum) much less perfectly known, inhabiting the upper.
It is possible that the latter as considered herein consists in reality of
several subspecies.

The species shows several characters that place it close to apama.
The principal one of these is the large fulvous or gray area on the fore
wing below. The chief constant difference between the two (apama
homoperplexa and dumetorum perplexa approach each other so closely
that aside from this basic character they are almost inseparable) lies
in the shape of the outer margin. In dumetorum it is convexly angled
in the vicinity of M2, and straight or slightly concave below it to the
inner margin, while in apama the whole margin is evenly convex.

220 bulletin: museum of comparative zoology

Callophrys dumetorum dumetorum Boisduval

Thecla dumetorum Boisduval, 1852, Ann. Soc. Ent. France (2) 10, p. 291;
Oberthur, 1913, Et. Lep. Comp. 9, p. 40, pi. 236, fig. 1926; Draudt, 1919,
in Seitz, Macrolep. World, 6, p. 763, pi. 154b; id., 1924, p. 1043. (all
partim)

Thecla dumetorum: auct. (partim)

Callophrys dumetorum: auct. (partim)

The type, in the United States National Museum, came from Cali-
fornia. No definite locality was designated. Due to a lack of material
agreeing satisfactorily with the description and with Oberthur's
figure of the type (loc. cit.) no type locality has been fixed.

Localities. California: Phelan; Snow Creek; Dana Point (all
H.K.C.); Pasadena (M.C.Z.); Calistoga; Petrified Forest (both in the
collection of R. G. Wind). These are all doubtfully typical dumetorum.

Male above uniform brownish gray. Fringe white, basally gray.
Female similar but more brownish, and with a discal fulvous suffusion.

Male below with fore wing gray or fulvous, inner margin somewhat
paler. Costa and outer margin as far down as Cui green. Hind wing
uniform green with two or three white spots, the principal ones being
on the costa and in the CU1-CU2 interspace. Fringe gray, slightly
paler outwardly and at the vein-ends on the hind wing. Female
similar, but with the gray area on the fore wing more fulvous. Both
sexes frequently have a more or less strongly developed discal line on
the fore wing, usually merely an intensification of the gray or fulvous.

Length of fore wing. Male, 12.5-13 mm.; female, 13 mm.

The above rather brief description is based on three males and a
female (two males from Phelan, one from Snow Creek, and a female
from Dana Point) which may or may not be typical dumetorum.
Several points in Boisduval's description differ from those found in
these specimens (namely, the fulvous (of Boisduval) instead of gray
color of the large area on the fore wing below; the presence of a number
of white spots on the hind wing below, lacking in the above-described
specimens). Typical dumetorum was very likely described from much
further north than these four specimens, although just where is still
unknown. The Calistoga specimens, while apparently approaching
Boisduval's description a little more closely than these, still differ
enough to be closer to the more southern specimens.

These specimens, of what is here taken to be typical dumetorum,
differ from perplexa in the absence of fulvous edging on the costa of the
fore wing below, and (in the male) by the gray, instead of fulvous,
area on this surface. There are, however, hints of fulvous on this area,

clench: notes on lycaenid butterflies 221

but this may possibly be explained by the fact that the specimens
came from a region not too remote from the home of true perplexa.

Two of the three ( lalistoga males examined have a definite fulvous
shading on the hind wing above, and one of them a patch on the fore
wing as well. More specimens are needed to tell whether or not this is
a local race of dumetorum. This character is quite unusual for dume-
torum and appears not to have been noticed previously.

Callophrys dcmetorum perplexa Barnes and Benjamin

Thecla affinis: auct. (partim)

Thecla dumetorum: auct. (partim)

Callophrys dumetorum: McDunnough, 1914, Ent. Rec. and Journ. Variation,

26, p. 196. (partim)
Callophrys dumetorum race perplexa Barnes and Benjamin, 1923, Contrib.

Nat. Hist. Lep. North America, 5, p. 65; Draudt, 1924, in Seitz, Macrolep.

World, 5, p. 1043. Comstock, 1927, Butterflies of California, p. 168,

pi. 50, figs. 17, 18, 19 (partim); Hoffman, 1940, Anales Inst. Biol. Mex., 11,

p. 708 (no. 623).

Holotype, allotype and a number of paratypes in the United States
National Museum. Described from San Diego, California.

Other localities. California : Pine Valley ; Corona ; Balboa ; El Modena
(all in coll. H.K.C.). Mexico: (northern localities, adjacent to Cali-
fornia).

Differs from typical dumetorum (as considered in this paper) in the
fulvous edging on the costa of the fore wing below, and in the in-
creased fulvous in the discal region of this same surface.

Apparently restricted to southern California and adjacent Mexico.

Length of fore wing. Male, 11-13.5 mm.; female, 11.5 mm.

Callophrys comstocki Henne
Callophrys comstocki Henne, 1940, Bull. So. Cal. Acad. Sci., 39, p. 71.

Holotype and allotype from the Providence Mts., San Bernardino
County, California, in the collection of the Los Angeles County
Museum. Paratypes from the same locality in the United States
National Museum, Canadian National Collection. Also (?) in the
collections of C. H. Ingham and C. Henne.

Topotypes have been examined from the collections of A. C.
Frederick, D. B. Stallings and the author.

Above, both sexes slate gray. Male with a small oval scent-pad.
Fringe gray, paler outwardly. Hind wing but slightly scalloped.

222 bulletin: museum of comparative zoology

Below, both sexes flat grayish green. Fore wing with inner marginal
area, centrally as far costad as M3, gray. This wing crossed by an
obsolescent post-discal line of white, basally gray-black, dashes.
Hind wing with a similar, but more distinct, line, edged basally with
black. This line outwardly displaced in M3-C111, and irregular thence
to the inner margin. Fringe of both wings basally gray, outwardly
white.

Length of fore wing. Male, 10-11 mm.

Closest to apama, but differing in several respects, principally the
gray upper surface of both sexes, and the flat green below, unrelieved
by any fulvous shading. It is a very interesting and isolated species,
possibly a local modification of apama, but certainly distinct enough
to warrant full specific distinction. So far as known it is restricted to
the Providence Mts.

Callophrys apama Edwards

This species occurs over a relatively compact range from Arizona
to New Mexico and north to Colorado, becoming differentiated in the
latter regions into a distinct subspecies. The limits of range of this
species are fairly well-known. It may, however, be turned up in
eastern California (doubtful) or southern Utah. (The Carnegie
Museum has a single male from this state; I have not seen it.)

From its near relative, C. dianetorum, apama may be distinguished
by the character given above under the general remarks for dume-
torum.

Another close ally (perhaps its closest) is the recently described
C. comstocki. This latter may be separated at once by its complete
lack of fulvous shading on the underside, by its smaller size, and by
the more distorted, less prominent white discal line on the hind wing
below.

Callophrys apama apama Edwards

Thecla apama Edwards, 1882, Papilio, 2, p. 137; Draudt, 1919, in Seitz, Macro-

lep. World, 5, p. 763; id., 1924, p. 1043.
Callophrys apama: Barnes and Benjamin, 1923, Contrib. Nat. Hist. Lep.

North America, 5, p. 67.

Type locality. Fort Grant in the Graham Mts., Arizona. One male
and two females labelled "Arizona" in the Edwards collection at the
Carnegie Museum. One of the latter made a lectotype by Holland.

Other localities. Arizona: Oak Creek Canyon (6000 ft.) (M.C.Z.);
White Mts.; Catalina Mts. (both in coll. H.K.C.); Navajo Mt. (in

clench: notes on lycaenid butterflies 223

coll. R, G. Wind); Sierra Aneha Mts. (in coll. P. S. and C. L. Reming-
ton).

Above in the male dark gray with a brownish tinge. A small,
almost circular scent-pad at the upper cell-end. Fringe of fore wing
concolorous with ground (or slightly darker), paler outwardly; of
hind wing similar, but frequently almost pure white outwardly; tips of
veins Cuu Cu2j and 2A usually white.

Female similar, but with fulvous patch on fore wing and usually
on hind wing. Fringe outwardly slightly paler.

Below, fore wing largely fulvous. Inner margin gray. A post-
discal, white irregular line crosses from costa to Cu2, composed of
white dashes, basally edged with black. Base of wing and apical area
outside of post-discal line (and extending down as far as Cux) green.
Hind wing green. A post-discal line, similar in construction to that of
the fore wing, but here more tortuous, crosses from costa to inner
margin, outwardly displaced often in the form of a crude "\Y" in the
M3-Q12 region. Basal to this line runs a closely adherent band of
fulvous. Occasionally there is a definite suggestion of a Cui-Cu2 sub-
marginal spot. Fringe of fore wing basally dull fulvous, outwardly
gray; of hind wing similar, but paler outwardly and tipped at Ciii, Cu2
and 2 A with white.

Length of fore wing. Male, 10.5-13 mm.; female, 10.5-11 mm.

The typical subspecies is distinguished from its Colorado repre-
sentative by the clarity and completeness of the markings below,
which are reduced frequently to the point of complete obscurity in
homoperplexa.

Callophrys apama homoperplexa Barnes and Benjamin

Thccla dumetorum auct. (partim)

Callophrys apama race homoperplexa Barnes and Benjamin, 1923, Contrib.

Nat. Hist. Lep. North America, 5, p. 68; Draudt, 1924, in Seitz, Macro-

lep. World, 5, p. 1043.

Holotype, allotype, and several paratypes in the United States
National Museum. Holotype and allotype from Denver, Colorado.
Paratypes from Golden, Boulder, and Denver, Colorado.

Other localities. Colorado: Durango; Husted and Starr Ranch
(El Paso County) (all M.C.Z.); Boulder Canyon; Eldora (both in
coll. P. S. and C. L. Remington). New Mexico: nr. Hot Springs, Las
Vegas (7000 ft.); Rincon (both in M.C.Z.). Specimens from Golden
have also been examined.

224 bulletin: museum of comparative zoology

On the upperside it differs apparently only in the more fulvous
shading of the female, and a greater tint of fulvous in the male.

On the underside the differences are much more pronounced.
Homopcrple.va lacks, in varying stages of completeness, the white
discal lines on each wing which form such a conspicuous part of typical
apama. The green on the fore wing appears to be more greatly ex-
tended.

Length of fore wing. As in the typical subspecies.

This subspecies is very interesting, and would seem to point to-
wards a connection with affinis. This at first implausible suggestion
becomes more likely when one notices that males of this subspecies are
frequently dull fulvous, and that occasionally specimens are found in
which the green of the underside of the fore wing covers an abnormally
large part of the wing, or shows itself faintly as a discal green suffusion.
Another point in favor of this alliance is the tendency of affinis to
produce specimens with hints (sometimes startlingly complete) of a
discal white line on the hind wing below. Yet another point is that the
ranges of the two do not apparently overlap, or if so, they only occur
sympatrically in a very restricted area. More material is needed from
the zone of transition (northern Colorado and adjacent regions) to
prove or disprove this.

Prior to the work of Barnes and Benjamin, this subspecies was com-
pletely confused with dumetorum perple.va (dumetorum auct.) of
southern California, and very understandably so. The two are strik-
ingly similar, and frequently almost the only real point of distinction
(superficial) is the difference in the fore wing outline. In general, how-
ever, males of homoperplexa have a more ruddy upper surface than do
males of perple.va. The green below of homoperplexa tends to be some-
what more brassy than that of perple.va.

Extreme homoperplexa lacks any suggestion of the discal line on
either wing below. Throughout Colorado, however, and particularly
in the southern part of the state specimens are frequently found that
tend quite definitely toward the typical subspecies. The two New
Mexico examples (localities given above) are in this category as well.
Apparently apama is restricted to Arizona.

Callophrys affinis Edwards

The exact range of this species is not yet known. It extends, from
the material and information at present available, from Utah north to
Wyoming and Washington. It may possibly occur in Colorado also.

clench: notes on lycaenid butterflies 225

The Washington specimens appear sufficiently distinct to warrant
separate racial consideration. Wyoming specimens appear to be
transitional in some respects, but are referred at present to the typical
subspecies.

This species is distinguished by its large, usually entirely unmarked
green under surface, and (with the exception of the Washington race)
large amount of fulvous above in the male. Both dumetorum (Calis-
toga) and apama homoperplexa have this suffusion in the males, but
not to the extent of typical affinis.

Callophrys affinis affinis Edwards

Theda affinis Edwards, 1862, Proc. Acad. Nat. Sri. Philadelphia, p. 223;

Draudt, 1919, in Seitz, Macrolep. World, 5, p. 763; id., 1924, p. 1043.
Callophrys affinis: Barnes and Benjamin, 1923, Contrib. Nat. Hist. Lep. North

America, 5, p. 66.

The types, one male and one female, from Utah, are in the Carnegie
Museum. Silver Lake, Utah, is here selected as the type locality,
based upon two males and two females in the M.C.Z. Barnes and
Benjamin (loc. cit.) referred to some specimens from Silver Lake as
topotypes, but did not elaborate.

Other localities. Wyoming: Teton Mts. (in coll. R. G. Wind).
Edwards' description of this species is very good, and is quoted here:
"Both sexes glossy red brown, brightest in female; the male has a
smooth oval spot on disc of primaries; costa of primaries and base of
both wings blackish brown; whole hind margin edged with same
color; fringe white.

"Under side uniform apple green, except on inner margin of pri-
maries, where it is pale, brownish grey; both wings immaculate; costal
edge of primaries grey; hind margin of secondaries without crena-
tions."

Length of fore wing: Male, 12-13 mm.; female, 12.5-13.5 mm.

The typical form is large and bright. It differs from washingtonia
in the shade of apple green below, and in the large extent of fulvous
on the male above.

Callophrys affinis washingtonia, new subspecies

Upperside:

Male. Both wings dark gray, slightly brownish, and occasionally
suffused with dull fulvous in the disk. Fringe white, basally dark
gray.

226 bulletin: museum of comparative zoology

Female. Similar to the male, but with the fulvous more extended.
Underside:

Male. Both wings green, slightly bluish, and very faintly brassy.
Inner margin of fore wing to 2A, and frequently to C112, gray. Hind
wing occasionally with the merest suggestion of a spot in the CU1-CU2
interspace.1 Fringe white, slightly darker at the vein-ends on the
hind wing.

Female. Similar to the male, but with the addition of a very thin,
extreme marginal line of white, running from anal angle to Mi. This
is very likely an individual variant, and will probably not be found to
hold true for other females. At the anal angle is a very small white
patch, the origin of the white line, and probably also merely an
individual variation.

Length of fore wing. Male, 11-12.5 mm.; female, 12 mm.

Holotype. Male, Alta Lake, Washington, April 25, 1935 (J. C.
Hopfinger) ex coll. P. S. and C. L. Remington.

Allotype. Female, same data as holotype.

Paratypes. Two males, Brewster, Washington, May 9, 1939 (J. C.
Hopfinger?), ex coll. L. P. Grey.

Holotype and allotype, no. 26259 in the M.C.Z. Paratypes in the
author's collection.

Remarks. Differs from typical affinis in the much more reduced
fulvous above in both sexes, and in the more bluish tone of the green
below.

A series of over 15 examples from the Teton Mts., Wyoming, seems
about intermediate between washingtonia and true affinis. There is,
however, little if any variation in the amount of fulvous on the fore
wing above in the male, while this character in washingtonia varies
from a limited amount (smaller than the smallest .of the Teton Mts.
specimens) to none at all. In several of the Wyoming specimens
there is an indication of a tendency to produce a row of white dashes
on the hind wing below; in one specimen quite strongly.

Callophrys viridis Edwards

Thecla viridis Edwards, 1862, Proc. Acad. Nat. Sci. Philadelphia, p. 223.

Thecla dumetorum: auct. {partim)

Callophrys dumetorum: Barnes and Benjamin, 1923, Contrib. Nat. Hist. Lep.
North America, 5, p. 64 {partim); Comstock, 1927, Butterflies of Cali-
fornia, p. 168, pi. 50, figs. 16, 20, 21, 25 (some of these are apparently
dumetorum) {partim).

1 The allotype has an indication of dashes in other neighboring interspaces, as well as a row
of very faint dashes on the fore wing. This latter can be seen faintly in certain lights in one
of the paratypes.

clench: notes on lycaenid butterflies 227

The type locality is here selected as San Francisco, California,
based on a female in the collection of P. S. and C. L. Remington
(May 8, 1934, Wm. Hovanitz collector) that agrees excellently with
the original description. In the absence of any type at the Carnegie
Museum this specimen is here made the neoholotype.

Other localities. "California" (M.C.Z.).

The male is above uniform gray. Fringe white, gray basally and at
the vein-ends of the hind wing. Female similar, but may be largely
suffused with fulvous, leaving a dark base, costa and outer margin on
the fore wing, and a dark base only on the hind wing.

Below in both sexes both wings are uniform pea-green. Fore wing
with inner marginal area to vein Cu2 gray, save on outer margin,
where the green extends down to 2A. Costa edged with fulvous. In
the disk is a faint row of dull white dashes, three in number in the
specimen examined, in interspaces M2-M3-Cu1-Cu2. On the hind wing
this row is continued, commencing at the outer angle and proceeding
to just basad of the anal lobe on the inner margin. The marks com-
posing this line are rather displaced and irregular in size, the one in
Cux-Cu2 being the largest. Those on the fore wing and one on the costa
of the hind wing are basally edged with dark brown or black. Fringe
of both wings white, rather dull, and basally slightly darkened.

Length of fore wing. Female, 13.5 mm. (neoholotype).

This species, heretofore considered synonymous with dumetorum,
is perfectly valid. Dumetorum, as described by Boisduval,1 has a
large fulvous area on the underside of the fore wing, lacking in viridis.
This character (the extent of the green on the underside of the fore
wing), which so far as known is very constant and subject to almost
no individual variation,2 is alone enough to raise Edwards' name from
the synonymy. Oberthur's figure of the type of dumetorum. (loc. cit.)
confirms Boisduval's description in this respect. True dumetorum has
been discussed further above.

Even with viridis and dumetorum separated, the picture is still any-
thing but clear. In California, apparently right along with viridis,
occurs a very green form with an immaculate underside. With only a
limited number of specimens available, it has been impossible to de-
termine whether this is a distinct species or merely an extreme of
viridis. Either is quite possible, although the latter is the more
probable.

1". . . et le disque des ailes superieures est beaucoup plus largement roussatre, ce qui fait
que le vert domine nioius." (reference under dumetorum).

2There is a certain amount of individual variation in this character in C. apama homopler-
plera, as has already been noted, but this is apparently an indication of transition towards
another species (affinis), and as such is excusable.

228 bulletin: museum of comparative zoology

Several specimens of Callophrys have been examined from the
collection of Mr. R. G. Wind, having been collected by him chiefly in
the Sierras. These, while apparently close to viridis differ in several
respects from it. These specimens, however, also differ considerably
inter se, so that in the absence of genitalic examination or larger series,
nothing further can be said.

Callophrys sheridanii Edwards

This species occupies a long, narrow, mountainous strip from
Cloudcroft, New Mexico north as far as Brewster, Washington. It is
differentiated into at least two subspecies, the typical occurring in
Wyoming, Colorado and probably New Mexico, while the subspecies
neoperplcxa ranges from Utah north to Washington. Specimens from
the latter locality do not seem exactly typical of neoperplcxa and
when more specimens are available they may be found to belong to a
distinct race.

The chief character whereby this species may be differentiated from
any other now known is the long, nearly straight (usually) white line
on the hind wing below, frequently basally bordered with black. It is
also one of the smaller species of the genus.

Callophrys sheridanii sheridanii Edwards

Thecla sheridanii Edwards, 1877, in Carpenter, Field and Forest, 3, p. 48

(lapsus calami).
Thecla sheridanii: Draudt, 1919 in Seitz, Macrolep. World, 5, p. 763; id., 1924,

p. 1043.
Callophrys sheridanii: Barnes and Benjamin, 1923, Contrib. Nat. Hist. Lep.

North America, 5, p. 66.

In the Carnegie Museum is one female labelled "Bighorn, Mont."
(fide Sweadner, in lift.) here selected as neoholotype. Holland's selec-
tion of a lectotype from Denver, Colorado is invalid as that is definitely
not the type locality.

Other localities. Wyoming: Teton Mts. and vie. (A.M.N.H.).
Colorado: Chimney Gulch (M.C.Z.); Ft, Collins; Denver (both
Barnes and Benjamin, p. 66); Eldora; Ceal Creek (both in coll. P. S.
and C. L. Remington). New Mexico: Cloudcroft (M.C.Z.).

Male above dark gray with a slight brownish tinge. A small oval
scent-pad at the upper cell-end. Fringes of both wings white, gray
basally. Female similar, but lacking the scent-pad.

Male below with both wings slightly brassy green, irrorated faintly,

clench: notes on lycaenid butterflies 229

especially on the hind wing, with obscure black scales. Fore wing with
inner margin gray-brown. A post-discal line crosses from costa to CU2
(occasionally with an inwardly dislocated extension in ( u2-2A), white,
and basally narrowly lined with black-brown. Hind wing with a
similar line, but heavier) and running from costa to inner margin in a
nearly straight line (frequently dislocated, but seldom if ever curved).
A small, white cell-end spot is occasionally present. Fringe as on
upper surface, but slightly greenish towards outer angle. Female as
in the male.

Length of fore wing. Male, 10-12 mm.; female, 11 mm.

The typical form differs from neoperplexa in the thicker white line
below, and the more prominent basal black edging to this line. Barnes
and Benjamin state that a specimen from Cloudcroft, New Mexico is
closer to the subspecies neoperplexa. This is decidedly queer, if so,
and is not borne out by the single specimen from that locality in the
M.C.Z., which, although not perfectly typical, is close enough to
belong here for the present.

Callophrys sheridanii neoperplexa Barnes and Benjamin

Callophrys sheridanii race neoperplexa Barnes and Benjamin, 1923, Contrib.
Nat. Hist. Lep. North America, 5, p. 67; Draudt, 1924, in Seitz, Macrolep.
World, 5, p. 1043.

Holotype and allotype from Eureka, Utah. Paratypes from Stock-
ton and Silver Lake, Utah. Holotype, allotype and 2 male, 1 female
paratypes in the United States National Museum.

Other localities. Montana : Polaris (H.K.C.). Washington : Brewster
(H.K.C.). In the M.C.Z. is a series from Silver Lake.

Its differences as compared with typical sheridanii have been
pointed out above. The Brewster, Washington, specimens do not
agree perfectly with Utah neoperplexa, and may ultimately be found
racially separable. They are very close, however, and for the present
will be left under that name. It is possible that some of the Sierran
material from California, mentioned under viridis, will prove to be
southern extensions of sheridanii stock, racially modified.

Length of fore wing. Male, 10-11 mm.; female, 11-11.5 mm.

b. THE ACASTE GROUP OF THE GENUS THECLA

The species treated here form a more-or-less closely interrelated
group isolated by Draudt (1919, in Seitz, Macrolep. World, 5, p. 762)
as the tailless section of his amyntor-group, of the genus Thccla. No

230 bulletin: museum of comparative zoology

better classification can be made with accuracy at present, since the
generic subdivision of the neotropical Theclinae is a task no one has
yet attempted with any degree of completeness1.

That the "tailless section" is here considered separately from the
"tailed section" is due principally to a very great dearth of material
of the latter. The two sections are apparently very closely allied, at
least in appearance2, and are separable only in minor characters.

The acaste group3 may be distinguished from the "tailed section"
in the following external particulars : The anal angle of the hind wing is
more produced, and the anal lobe is longer, more prominent. In
general the tail is absent (see footnote no. 3) but if it is present it is
coarser (broader) and proportionately shorter than those of the other
section. Also on the hind wing, the vein-ends are here slightly more
tufted. The green below is less uniform and usually less shining. The
general appearance of this group is more reminiscent of CaUophrys, or
even, somehow, of Incisalia, while the others seem more typically
Thecline.

The various members of the acaste group are of average Thecline
size, ranging (in the length of the fore wing) from 12 (female of remits)
to 16 mm. .(female of a. acaste). The males all have scent pads4, but
they are usually small and almost unnoticeable, occasionally being of
the same color as the ground color, a rather unusual occurrence. The
males are above (excepting marialis) some shade of blue or purple,
very metallic, with dark borders of width varying with the species.
The females on this surface are duller, the blue being considerably
less metallic, and more basally restricted. Below the sexes are similar,
with a rather similar basic pattern : ground color green ; a post-discal
line (outwardly white, basally dark) on both wings; a submarginal
row of red spots on the hind wing; a marginal row of spots or patches,
or a marginal band, on both wings; a basal quadrate patch of dark
brown on the hind wing. Any or all of these may be suppressed or

:The members of this section (perhaps even more sc than those of the "tailed section") seem
to bear affinity to the genus CaUophrys Billberg. Indeed. W. D. Field (1939, Univ. Kansas
Sci nee Bull., 26, p. 347) has placed Thecla hercdotus Fabr. (a member of the "tailed section")
in this genus, without comment however.

2Due to the briefness of time available for study it was not possible to make any genitalic
preparations. These, when examined, may show further, more real differences.

3So called in preference to the "tailless section" of Draudt, since that method of division has
proved incorrect. Thecla longula (pastor) and Thecla marialis, both tailed species, are clearly
members of the group now under consideration, while a tailless species, quite obviously belong-
ing to the other section, is being described as new in another paper.

4Godman and Salvin (1887, Biol. Centr.-Am. Lep., Rhop. 2, p. 34) erroneously characterize
agricolor as lacking a scent pad.

clench: notes on lycaenid butterflies 231

variously developed. An inner marginal hand- of tan or gray is always
present on the underside of the fore wing.

The several species together occupy a wide range from Mexico to
Bolivia and southern Brazil, and even Argentina. Our knowledge of
the distribution of the individual species is very imperfect, and will
probably remain so for some time to come.

Key to species

This key was very difficult to compose, and in parts may be in-
accurate; it should be used, therefore, with this in mind. In several
of the species but one sex is known, thus making it impossible to in-
clude characters which are definitely known to apply to both male
and female of such species.

1 . Frons brown 2

- Frons green 7

2. Outer margin of hind wing below with red-gray edging, either as a definite

band, or as a series of internervular spots 3

- This margin without red-gray spots, being green, as in the rest of the

wing 6

3. Outer margin of fore wing below with red-gray edging; a well-marked

discal line as well on this surface a. agricolor (1)

- This margin without red-gray edging; discal line obsolescent or wanting . .4

4. Tail at Cu2 on secondary longula (5)

- No tail at Cu2 5

5. Small, expanse less than 30 mm. (usually about 26 mm.) remits (2)

- Larger, over 30 mm.; up to 32 mm. or so agricolor banosensis (la)

6. Male with scent pad black; underside nearly uniform green . .longuloides (4)

- Male with scent pad colored blue as in the ground color; below with bands

of darker green pseudolongula (3)

7. Upperside of male brown; tailed at Cu2 on secondary marialis (9)

- This surface of male bright blue; no tail at C112 8

8. White discal line on underside of hind wing (may tend toward obsolescence)

- No white discal line on this wing legionis (6)

9. White patch in center of gray area on underside of fore wing (only in

male?) portoena (8)

- No white patch in this area, male or female 10

10. Submarginal red spots on underside of hind wing absent (almost or com-
pletely); white discal line prominent acaste catharinetisis (7a)

— Submarginal red spots on underside of hind wing usually well-developed;
discal white line obsolescent a. acaste (7)

232 bulletin: museum of comparative zoology

1. Thecla agricolor agricolor (Butler and Druce)

Strymon agricolor Butler and Druce, 1872, Cist. Ent., 1, p. 105; Butler, 1873,

Lep. Exot., p. 158, pi. 57, fig. 4.
Thecla agricolor: Hewitson, 1877, 111. Diurn. Lep. Lycaenidae, p. 201; Godman

and Salvin, 1887, Biol. Centr.-Am. Lep. Rhop., 2, p. 34, pi. 52, figs. 11,

12; Draudt, 1919, in Seitz, Macrolep. World, 5, p. 762, pi. 154a; Hoffman,

1940, An. Inst, Biol. Mex., 11, p. 707.

The type is in the British Museum, presumably. It was taken in
Cartago, Costa Rica. Draudt gives the range as "Mexico to Panama."
Godman and Salvin list Jalapa, Mexico; Duenas, Guatemala; Irazu
and Rio Sucio1, Costa Rica; Bugaba and Chiriqui, Panama. Hoffman
gives "Tierra templada de Vera Cruz. Sur de Puebla. Morelos.
Valle de Mexico (2250 M.)." Specimens in the M.C.Z. are all from
Jalapa, Mexico.

Above, in the male, metallic blue, duller than in the other species.
Both wings with a rather indefinite and heavy dark border on the outer
margin. Costa of fore wing narrowly black. Costa and inner margin
of hind wing narrowly gray. Anal lobe of hind wing rusty, black-
fringed. The scent-pad is very small, almost unnoticeable (see foot-
note, p. 230), and lies just beyond the upper cell-end.

Below, fore wing gray from inner margin to M3, darker adjacent
to the cell. Remaining area green. Outer margin from apex to inner
angle with a moderately heavy grayish border. A discal line of
reddish crosses from costa, disappearing shortly below M3. A faint
submarginal line behaves similarly. Hind wing with a broad marginal
border, outwardly gray, basally reddish. The basal limit of this border
is, in the M3-CU1 and 2A-inner margin interspaces, orange. A discal
line of dark green and reddish scales crosses the wing obscurely. Base
marked with a large, quadrate, almost black patch. Anal lobe maroon.

The female is duller above than the male, with the blue more re-
stricted, leaving very broad costal and outer marginal borders on the
fore wing, and a broad outer marginal border on the hind wing. It is
almost exactly similar on this surface to pseudolongula, but the outer
limit of the blue is more sharply defined.

Below, the female is similar to the male, but with the markings
slightly brighter.

Length of fore wing. Male, 12.5-13.5 mm.; female, 14.5 mm.

'Possibly this refers to a Rio Sucio in northern Colombia, emptying; into the Gulf of Darien
just below the south-eastern tip of Panama. The specimens on which this record is based may.
therefore, be transitional to bahosensis.

clench: notes on lycaenid butterflies 233

la. Thecla agricolor banosensis, new subspecies

Upperside:

Female. Both wings bright metallic blue. Fore wing with a costal
and outer marginal border of black-brown, covering the whole cell-end-
to-apex region, and narrowing towards the inner angle. Hind wing
with a gray costal border, becoming black-brown on the outer angle
and outer margin. Inner margin gray. Anal lobe rusty, the color
extending basad slightly, and costad as far as Cu2. Fringe of both
wings black-brown, paler between the veins.
Underside:

Female. Both wings green. Fore wing with a paler, apple-green
marginal stripe and a discal row of three obscure red-brown spots
(Mi-M2-M3-Cui). Inner margin to Cu2 gray, black basad, next the
cell. Between the gray and the green, in the Cui-Cu2 interspace,
is a band of fulvous. Hind wing with a small black basal patch, a
discal transverse line of gray, thin costad of Cui, heavier thence to
inner margin. A submarginal row of obsolescent red lunules from costa
to inner margin, replaced costally by dark green. On the outer margin
is a band of hoary maroon, basally scalloped from Mi to the anal
angle. Anal lobe maroon. Fringe of both wings white-gray, darker
at the vein-ends.

Length of fore wing. Female, 15 mm.

Holotype. Female, San Pablo, Rio Pastaza, vie. Banos, Ecuador,
2200 meters(?), (Clark-Maclntyre), ex coll. F. M. Brown, in the
American Museum of Natural History.

Remarks. This subspecies differs from typical agricolor1 in several
respects, namely : the absence of the marginal hoary band on the fore
wing below, reduction in size of the black basal patch of the hind wing
below, and the reduced size of the marginal band of the hind wing
below, and its differentiation into two bands. The blue color above is
very bright, far brighter than in any other female of this group thus
far examined (except for a single female of pseudolongula, of almost
the same intensity). This character, however, is in all likelihood of
no significance, being merely an age factor.

This subspecies, with its reduced markings below, strongly sug-
gests a transition from agricolor to rcmus, which, in turn, may connect
to pseudolongula. Many more specimens are needed, however, before

'The Jalapa, Mexico, specimens and the holotype of bafwsensis were compared with Butler's
figure in the Cist. Eat. (I.e.) and the former agreed almost perfectly. They were therefore used
in the following comparison as typical agricolor, even though not topotypical.

234 bulletin: museum of compaeative zoology

this suggestion can be proven or denied. Genitalic examination would
help considerably.

2. Thecla remus Hewitson

Thecla remus Hewitson, 186S, Descr. Lycaenidae, p. 34; 1877, 111. Diurn. Lep.

Lycaetiidae, p. 201, pi. 80, figs. 655-656; Draudt, 1919, in Seitz, Macrolep.

World, 5, p. 763, pi. 154b.
Thecla deidamia Burmeister, 1879, Atlas de la Descr. Phys. Rep. Argentine, 5,

pt. 2, p. 24.

Described from Brazil. Type (female) is presumably in the British
Museum, although Mr. Goodson (who examined the British Museum's
specimens of the present group for me) wasn't exactly clear on that
point in his notes. Draudt in Seitz gives no additional information,
and does not seem to have known the species. Burmeister records it
(as deidamia) from Las Conchas, north of Buenos Aires, Argentina.
Four specimens, all females, in the M.C.Z.: three from Blumenau,
Sta. Catharina, Brazil, and one, ex coll. J. Doll, labelled "Brazilia."
One male, in the American Museum of Natural History from "Massa-
randuba-Blumenau, Brazil."

Male above brilliant iridescent blue, violet-tinted in some lights.
Both wings edged with black on the outer margins. Hind wing gray
on costa and inner margin. Anal lobe dull maroon.

Below, the male is green on both wings. Fore wing with gray on
inner margin to Cu2, darkened basally. Hind wing with a marginal
row of hoary, reddish-gray spots, almost connected. A discal row of
obsolescent, irregular spots crosses the wing, between which and the
marginal row of spots is a suggestion of a row of red dashes, in M3-
CU1-C112. Anal lobe dark maroon.

The female above with the blue much duller and more restricted;
quite similar in appearance to the females of most of the other species
of the group. Anal lobe rusty. Below green, with the inner margin of
the fore wing gray, darker basad. Hind wing on the outer margin
with a series of internervular, almost round, reddish gray spots. This
wing crossed in the disk by a rather tortuous line of white, interrupted
frequently, and of varying intensity, basally edged with blackish.
Between this line and the marginal spots are one or two, rarely more,
red dashes, slightly crescentiform. Anal lobe dark maroon. Hewit-
son's descriptions (1868, 1877) of the female tallies quite well with what
is here regarded as remus, save for a few minor differences. The fulvous
patch at the outer angle of the hind wing below mentioned by Hewit-

clench: notes on lycaenid butterflies 235

son is very likely the result of wear. He mentions in both descriptions
a band of three or four spots on the fore wing below, near the eosta.
No indication of such spots was found on the four specimens examined.
He does not mention the subminimal scries of three or four crescenti-
form dashes on the hind wing. This character, however, is apparently
variable, being lacking in one of the four M.( \Z. specimens, and almost
lacking in another. On the strength of these apparent differences it
was decided not to select a type locality for remus until either speci-
mens are found that match more closely the description, or it is
proven that he had a slightly aberrant example.

The male agrees quite well with Hewitson's 1877 description and
figure, save for the "green tint" lie speaks of. This, however, is fre-
quently caused by chemical action, and can be overlooked. It is above
close to males of pseudolongula, but may be separated by the broader,
more apically thickened outer marginal border.

Thecla deidamia is a name that ever since its publication in 1S79
appears to have been almost overlooked. Seitz made no mention of it,
and the only published reference of it known to the author is that of
Weeks (see synonymy under portocna).

Weeks referred his name to "Ruschew.", but this is in error, as a
glance at the original description will show. Ruscheweyh collected the
specimens, noted that they were probably new, and suggested the
name deidamia (in Hit to Burmeister, apparently). Burmeister, how-
ever, wrote the description and applied the name to it, wherefore it
must go to him.

With regard to the present placing of the name, it can be said only
that a comparison of Burmeister's description with Hewitson's
descriptions and figures (1868 and 1877) and several specimens of
remus revealed an almost perfect resemblance. Burmeister's locality
(Las Conchas, nr. Buenos Aires, Argentina) is quite in accord with our
present knowledge of the distribution of remus, although extending
its range somewhat southward.

3. Thecla pseudolongula, new species

Thecla longula: Hewitson, 1877, 111. Diurn. Lep. Lycaenidae, p. 200, pi. 80,
figs. 651, 652, 653, 654; Draudt, 1919, in Seitz, Macrolep. World, 5, p. 762,
pi. 154a. {nee T. longula Hewitson, 1868 (q.v.)).

Eyes narrowly ringed with white. Frons with two parallel rows of
long, partially erect, dark brown hairs, flanked outwardly (along the
rims of the eyes) with a row of rusty scales on each side and a band of

236 bulletin: museum of comparative zoology

shorter hairs, with rusty ones intermingled, between them. Caudad of
the antennae is a transverse row of long rusty hairs. Collar of long
hairs, bluish and rusty on top, becoming intermingled with white on
the sides. Palpi rusty outwardly, bluish white within; terminal joint
completely rusty. Antennae black above, white annulate below; club
black, tipped with fulvous and paler below (becoming white basad).
Thorax of the male above metallic blue or green, overlaid moderately
with long, anally directed hairs, heaviest next the abdomen; female
paler, more steely blue above; male below covered with dense brown
hair, paler in the female. Abdomen of the male above metallic blue,
very bright; belbw, yellow; tip above and below gray: of the female,
duller, the blue more steely and more anteriorly restricted; below
similar. Legs (absent in most of the specimens examined) apparently
brown, tarsi black and white annulate.
Upperside :

Male. Bright metallic blue, greenish in some lights, purplish in
others. The scent pad is so small as to be almost unnoticeable, con-
sisting merely of a short row of scales along the upper disco-cellular
at the cell-end. A marginal border, about 1 mm. thick, edges both
wings. On the fore wing it is slightly thicker than on the hind wing,
and towards the apex it expands still more (2-3 mm.). The anal lobe
is prominent and rust-colored. Fringe brown on fore wing; dull white
on hind wing, basally and at the vein-ends darker.

Female. Dull gray-blue, with very broad blackish borders, some-
what narrower and basally crenulate on the hind wing. Anal lobe as in
the male. Fringe also as in the male.
Underside:

Male. Both wings bright emerald-green, with the inner margin of
the fore wing broadly gray. The hind wing is marked by two trans-
verse bands of lighter green, rather indistinct, basad to the inner of
which is a suggestion (consisting usually of one or two obscure points)
of a post-discal line. In the extreme base is a smallish dark brown area.
Anal lobe rusty and adjoining it in the CU2-2A interspace is an obscure
rusty -hoary patch.

Female. Similar, but the markings more distinct.
Length of fore wing. Male, 13.5-15 mm.; female, 14-14.5 mm.
Holotypc. Male, Mapoto, Ecuador, ex A. G. Weeks collection.
Allotype. Female, no locality, ex R. M. Gray collection.
Paratypes. On male, R. Guamlo (or Guamba- label poorly written),
Ecuador, ex A. G. Weeks collection; one male, "Colombia", Oct. 10,
1913, ex F. A. Eddy collection; two males, no locality (possibly

clench: notes on lycaenid butterflies 237

Bogota, Colombia), ex A. G. Weeks collection; one female, no Locality
("So. Am."), ex C. J. Paine collection; two males, one female, vie.
Banos, Ecuador (Clark-Maelntyre), as follows: one male, Tunguragua,
1900 meters, March, 1939; one male, Runtim, 2000-2500 meters,
Nov. 26, 1938; one female, Rio Blanco, 1700-1900 meters, Oct. 19,
1938.

Holotype, allotype, and three male paratypes, no. 26223 in the
Museum of Comparative Zoology. One male and one female paratype
in the author's collection. The last three paratypes in the collection of
the American Museum of Natural History.

Remarks. Quite different from true longula, for comparison with
which, see under that species. The closest ally of pseudolongula yet
discovered appears to be longuloides, from which it differs in the
smaller scent-pad, broader borders and larger anal lobe. It is also
allied to remits, but that species has a marginal row of hoary spots on
the hind wing below.

4. Thecla longuloides, new species

Eyes hairy, narrowly ringed with white. Frons consisting of two
parallel rows of long, erect, dark brownish hairs, thickly intermingled
with rusty ones. Just outside these rows, paralleling the white eye-
margin, is a row of rusty scales. Basad of the antennae is a transverse
row of rusty hairs, and between the antennae a few white ones.
Collar of long, rusty and brown-black hairs. Palpi outwardly covered
with mingled pale blue, rusty and gray scales, inwardly almost en-
tirely pale-blue. Antennae black above, white annulate below; club
black above, below tipped with dull fulvous, backed by white. Thorax
of the male above metallic greenish blue, covered, chiefly on the
periphery (behind the head, along the sides above the wing bases and
anterior to the abdomen), with long, grayish-rusty hairs; thorax of the
female grayer blue above, without the greenish tinge: below tufted
with brown in the male, paler in the female. Abdomen of the male
above brilliant metallic green, below yellow; tip gray above and
below; abdomen of the female with the blue more anteriorly restricted
above (the remaining area gray) ; below as in the male. Legs black
and white annulate.
Upperside:

Male. Both wings brilliant metallic blue, greenish in some lights
and purplish in others. Fore vying with an almost linear black scent-
pad lying along the upper discocellular at the cell-end. Costa very

238 bulletin: museum of comparative zoology

narrowly, outer margin slightly more broadly black-bordered. The
latter thickens apically and extends briefly basad on each vein. Hind
wing with costa and inner margin gray. Outer margin narrowly black,
also extending basad for a short distance on the veins. Anal lobe small,
rusty colored. Fringe of fore wing brown, paler outwardly; of hind
wing similar, but whitish outwardly between the veins.

Female. Both wings dark gray-brown, costa and inner margin of
hind wing pale brown. Fore wing with dull steely blue from inner
margin to upper discocellular, from base to cell-end, outwardly down
to inner margin three-quarters out. Hind wing similar, leaving only
a narrow dark border on the outer margin, hazy and indistinct basad,
that thickens slightly towards the outer angle. Anal lobe as in the
male. Fringe as in the male, but darker.
Underside:

Male. Both wings uniform green. Fore wing with the inner margin
to just over Cua gray, becoming sharply black along the lower disco-
cellular. Hind wing with two very faint and obscure small white spots,
each lined basally with a few red scales: one post-discal in the CU1-CU2
interspace; the other submarginal in the 2A-3A interspace, touching
2A. The faintest indication of a submarginal band, almost un-
noticeable, consists merely of a very slight darkening of the green.
Fringe as on upper surface. Anal lobe obscurely dark rusty colored.

Female. Similar to the male, but lacking the black along the lower
discocellular of the fore wing, and with the faint submarginal band of
the hind wing a little more prominent. The green on the outer margin
of the fore wing extends down more into the (A12-IA interspace.

Length of fore wing. Male, 14 mm.; female, 13 mm.

Holoti/pe. Male, Coroico, Bolivia, May, 1899 (Wm. J. Gerhard),
ex A. G. Weeks collection.

Allotype. Female, Chulumani, Bolivia, Dec. 12, 1898 (Wm. J.
Gerhard), ex A. G. Weeks collection.

Holotype and allotype, no. 26224 in the Museum of Comparative
Zoology.

Remarks. This species is allied to pseudolongula, but may be dis-
tinguished from it, in the male, by the much narrower outer marginal
border above, and the presence of a larger, more definite scent-pad.
Both sexes have a considerably reduced anal lobe on the hind wing
(less than half the size of that occurring on pseudolongula). Below the
green is more uniform than in pseudolongula, with the light and dark
transverse bands almost non-existent. In the female the outer margin
of the fore wing does not seem to be so convex as in that sex of pscudo-

clench: notes on lycaened butterflies _'■>'>

languid, and the blue appears to be duller, although of about equal
extent.

5. Thecla longula Hewitson

Thecla longula Hewitson, 1868, Descr. Lycaenidae, p. 34. (nee longulti, Hewit-
son, 1877, and others).

Strymon pastor Butler and Druce, 1872, Cist. Ent., 1, p. 105; Butler, 1873,
Lep. Exot., p. 157, pi. 57, fig. 5; McDunnough, 1938, Mem. S. Cal. Acad.
Sci., 1, p. 24 (no. 364).

Thecla pastor: Godman and Salvin, 1887, Biol. Centr. Am. Lep. Rhop., 2, p. 34,
pi. 52, figs. 8, 9, 10; Draudt, 1919, in Seitz, Macrolep. World, 5, p. 762,
pi. 154a; Hoffman, 1940, An. Inst, Biol. Mex., 11, p. 707.

The type apparently is not in the British Museum. Mr. F. W.
Goodson, at Tring, informs me through Dr. Riley that there is no
specimen of longula (by which he meant pseudolongula, probably
exclusively) in the British Museum from Central America. The four
specimens of ''longula" from the Hewitson collection are all referable
to pseudolongula.

Five specimens of longula have been examined and compared with
the description. They seem typical, and in the absence of a type are
made neotypes, as follows:

Xcoholotypc. Male, Orizaba, Mexico, June 1941 (Stallings and
Turner).

Neoallotype. Female, same data as above.

Neoparatypes. One female, same data as above; two females,
Presidio, Mexico, June 1941 (Stallings and Turner).

Xeoholotype and Neoallotype deposited in the Museum of Com-
parative Zoology. One neoparatype in the collection of the author.
The remaining returned to the collection of Mr. D. B. Stallings and
Dr. J. R. Turner.

This species has been subjected to a rather peculiar misidentifica-
tion, for almost the whole of its existence in the literature to date.
Hewitson's original description of longula is of a totally different
insect from that of his description and figure of 1877 (in the "Illustra-
tions"), as can be seen from the following extract from it (1868):
"Underside dull green, tinted with orange at the apex of the anterior
wing1. Posterior wing without tails:2 crossed beyond the middle by

UVohably due to wear: see under remus.

2Evidently variable in this species. One of Godman and Salvia's illustrations of pastor
showed tailless, and one of the type ineotype< serii>s of bmijula is also naturally without tails.
Possibly the species is in a state of either losing or acquiring them.

240 bulletin: museum of comparative zoology

two bands of indistinct distant red-brown spots : a series of marginal
red-brown spots, irrorated with white: the lobe red-brown." Compar-
ing this description with that of his published 1877 (pseudolongida of
this paper) shows immediately that the two are not conspecific. When
the above description was checked against pastor, the true identity
of the name was shown.

6. Thecla legionis, new species

Eyes hairy, ringed with white or pale green. Frons green. Collar
and palpi in the present specimen indiscernible. Antennae black and
white annulate, nearly all black above; club black, tipped with fulvous,
more extensive and white-backed below.- Thorax pale steely blue
above, fulvous beneath, rather pallid where visible. Abdomen gray
above (pale bluish basad), cream below.
Upperside:

Female. Both wings steely blue. Fore wing broadly black on costa,
apex and outer margin. Hing wing more narrowly so on outer margin.
Costa and inner margin gray. Anal lobe rusty. Fringe of fore wing
blackish, slightly paler outwardly; of hind wing sordid white, darker
at the vein-ends.
Underside:

Female. Both wings green. Fore wing with inner margin from Cu2
gray. Cu2 and outer margin along this gray are fulvous. Base of gray,
adjoining cell, darker. Hind wing unmarked save by a series of
obscure reddish dashes in the M2-2A interspaces slightly basad of the
submarginal area, and two almost unnoticeable white costal spots,
one each in the Sc-Rs-Mi interspaces. Outward of the reddish dashes
the green seems a little paler. Fringe of fore wing fulvous, outwardly
obscured by gray. Of hind wing fulvous, outwardly whitish, darker at
the vein-ends.

Length of fore wing. Female, 12.5 mm.

Holotypc. Female, Blumenau, Sta. Catharina, Brazil (B. Pohl),
no. 26225 in the collection of the M.C.Z.

Remarks. This species, unfortunately represented by but a single
female, appears to stand closest to acaste. It has the green frons of
acaste, but the almost unproduced anal angle of, for example, remus.
Below it looks somewhat similar to a small acaste without a discal
white line on the hind wing. Above, the blue is slightly paler than in
females of acaste, but this may be due to fading. Below legionis
differs from typical acaste (which subspecies it most closely resembles)

clench: notes on lycaenid butterflies 241

in the almost complete absence of a discal white line, the only indica-
tion being two almost costal white spots, so faint as to be hardly
discernible. The fringe is also more fulvous than in either subspecies of
acaste. The fulvous edging of the inner marginal area of the fore wing
below is also absent in both acaste and catharinensis, but this character,
like the fulvous patches mentioned by Hewitson in the descriptions of
longula and remus, may be due to Avear.

7. Thecla acaste acaste Prittwitz

Thecla acaste Prittwitz, 1865, Stett. Ent. Zeit,, 26, p. 31S; Draudt, 1919, in

Seitz, Macrolep. World, 5, p. 763, pi. 154a.
Thecla lycimna Hewitson, 1868, Descr. Lycaenidae, p. 33; 1877, 111. Diurn.

Lep. Lycaenidae, p. 203, pi. 80, figs. 663, 664, 665.

The type locality of acaste is Corcovado, near Rio de Janeiro,
Brazil. Under his lycimna Hewitson (1S77) merely gives "Brazil."1
Mr. Goodson has examined the Hewitson type for me, and it is ap-
parently quite in accordance with Prittwitz' original description, and
has the characters here used to separate true acaste from catharinensis.
Rio de Janeiro, Brazil, is here selected as the type locality of lycimna,
thus better insuring its permanent synonymy. The type of acaste,
Mr. Goodson suggests, is probably either in Munich, Berlin, or Greis-
swald, granting the collections at those localities to be still intact.

Draudt gives as records: Sao Paulo, Santa Catharina and Rio
Grande do Sul. The last two will probably refer to catharinensis.
Seven examples in the M.C.Z., Rio de Janeiro and Canto Gallo, Brazil.
The range of this, the typical subspecies, appears to be quite limited.

The male above is generally similar to longuloides, but with the
scent-pad concolorous with the ground. The ground color is slightly
duller than in longuloides, especially marginally, and the outer margin
is more broadly black. Below it is uniform green, with a gray inner
margin on the fore wing, an obsolescent white transverse line on the
hind wing running from costa to inner margin, outward of which is a
row of tiny bright red spots or dashes, the most prominent in Cui-Cu2.

Female above similar to the female of pseudolongula, but with the
outer limits of the blue less definite. Below as in the male.

Both sexes differ greatly from pseudolongula and longuloides in the
possession of a green frons.

Length of fore wing. Male, 15-15.5 mm.; female, 14-16 mm.

!The original description (1868) cited no locality whatsoever.

242 bulletin: museum of comparative zoology

7a. Thecla acaste catharinensis, new subspecies
Thecla acaste: Draudt, 1919, in Seitz, Macrolep. World, 5, p. 763 (partim.

Upperside:

Male. Both wings dully shining violet blue, deepening slightly
towards the margin. Outer margin of both wings narrowly black,
thickening slightly towards the apex on the fore wing. Costa of fore
wing also narrowly black. Costa and inner margin of hind wing gray,
the latter shaded basally with bluish. On the hind wing the anal lobe
is rusty, fringed with black.

Female. Both wings black-brown with the basal two-thirds of the
fore wing and the majority of the hind wing dull steely blue, leaving a
costal, broad apical, narrower outer marginal dark border on the fore
wing, and a gray costa and inner margin on the hind wing, with a still
narrower dark outer marginal border. Anal lobe as in the male.
Underside:

Male. Both wings bright green. Fore wing inner margin gray to
Cu2, the green encroaching only at the outer margin. Base of this
gray area dark along the lower Dc. Hind wing with a prominent,
mildly tortuous white line, basally and obscurely bordered with red.
This line commences two-thirds out on the costa and proceeds nearly
straight to three-quarters out on inner margin. Anal lobe black,
shading into deep red basad. A few minute black scales in the sub-
marginal area.

Female. Similar to the male.

Length of fore wing. Male and female, J4 mm.

Holotype. Male, Santa Catharina, Brazil, ex A. G. Weeks collection.

Allotype. Female, Blumenau, Sta. Catharina, Brazil (B. Pohl).

Paratype. One female, "Brazilia", ex J. Doll Collection.

Holotype and allotype, M.C.Z. no. 26226. Paratype in the collec-
tion of the author.

Remarks. Differs from typical acaste in the more prominent and
complete white line on the under surface of the secondary, in the
absence of the submarginal red dots or dashes on the same wing below,
and, in the male, by the reduced marginal border above.

8. Thecla portoena, new species

Thecla deidamia: Weeks, 1905, 111. Diurn. Lep. (111. Unfig'd. Lep.), 1, p. 19.
(nee deidamia Burmeister: see under remits. The identity of the insect
Weeks called deidamia (loc. cit.) is determined by three specimens so
labelled in his collection, which are now made the types of portoena.)

clench: notes on lycaenid butterflies 243

Eyes hairy, ringed with white. Frons green. Collar dark rusty
above, shading to whitish below. Palpi rusty pale gray, dorsally
dark brown. Antennae black, white annulate below, and very faintly
above; club black, tipped with dull fulvous. Thorax dull black,
covered with bluish hairs, lightly on top, heavier laterally and next
the abdomen; below grayish tan. Abdomen above blue next the
thorax, gray thence to tip; below yellow, gray at the tip. Legs black
and white annulate.
Upperside:

Male. Both wings shining lavender blue, deepening towards the
margin. Fore wing with a rather broad, dark brown marginal border,
thickening apically. Costa with a similar, but narrower border.
Hind wing with costa pale gray. Inner margin gray, overlaid basad
with bluish scales. Basal area of wing overlaid with pale scales, giving
a rather hoary appearance to this region. Outer margin narrowly
black-brown, extending slightly basad on the veins. Anal lobe rusty,
black-fringed. Fringe of fore wing dark basally, paler outwardly; of
hind wing dark, white outwardly between the veins.
Underside:

Male. Both wings uniform green. Fore icing with the inner margin
gray-tan, with a white patch on the center. The green encroaches on
this gray-tan at the outer margin. Base of this area somewhat dark-
ened. Hind wing with a nearly straight white discal line, basally
edged with a few red scales, that runs from the costa towards the
inner margin, but disappears at about M3 or Cui. A submarginal series
of thin red dashes occupies the M3-2 A interpaces, and occasionally even
further costad. Anal lobe deep red, almost maroon, extending slightly,
in the form of a compact small area of reddish irroration, into the
Cu2-2A interspace.

Length of fore wing. Male, 12-13.5 mm.

Holotype. Male, Cusilluni, Bolivia, May, 1899 (Wm. J. Gerhard),
ex A. G. Weeks collection.

Paratypes. Two males: one male, same data as holotype; one male,
Coroico, Bolivia, April 20, 1S99 (Wm. J. Gerhard), ex A. G. Weeks
collection.

Holotype and one paratype, M.C.Z. no. 26227. One paratype in
the author's collection.

Remarks. T. portocna may be distinguished from both typical
acaste and its subspecies cathariensis as follows : the marginal border of
the fore wing above (male) is broader than in either; below, the white
line on the hind wing disappears before reaching the inner margin,

244 bulletin: museum of comparative zoology

while in both acaste and a. catharinensis it proceeds all the way; the
anal lobe is here smaller than in either, and colored deep red, while in
acaste and its subspecies it is black; the inner marginal area of the fore
wing below is here gray-tan, and has a central white patch, while in
acaste and catharinensis it is gray and has no such patch (in fact
portoena is the only species at present known to the writer that
possesses such a peculiar pattern character) ; T. portoena has the red
submarginal spots as in acaste (s.s.), but they are more linear, and seem
to extend further cost ad.

It is possible that portoena may be only a subspecies of acaste. The
differences between them, however, are such that two full species seem
involved, and while at present portoena seems to be a local representa-
tive of acaste, further knowledge of the distributions of the involved
forms may prove otherwise.

9. Thecla marialis, new species

Eyes ringed with pale green. Frons green. Palpi fulvous, scaled
outwardly with green ; terminal point black. Collar above green, with
pale greenish and fulvous hairs, shading to sordid gray on the sides.
Thorax above black with dull greenish-gray hairs frontad, laterally,
and next the abdomen, all back-directed; below covered with pale
rusty long hairs. Abdomen black-brown above, yellow below, dark
gray at the tip. Legs gray, with pale annulations.
Upperside:

Male. Both wings black-brown. Fore wing with the basal area over-
laid with dull olive-green, extending marginad roughly two-thirds.
On the upper cell-end is an elongated black scent-pad, rather small.
Hind wing similar but with the olive-green shading restricted more
basad. Anal lobe fulvous, this color extending basad along inner
margin for one-third its length, and along outer margin almost to Cu2.
Tail at Cu2, very short, but definitely present. Fringe of both wings
gray with a brownish tinge.
Underside :

Male. Both wings emerald-green. Fore wing with inner margin to
Cu2 gray, darkening to almost black, next the cell, and to darker gray
on the outer margin. Hind wing with an almost non-existent indication
of a post-discal line, the only real relic being a tiny white spot in
Cui-Ci^, capped by a minute red bar. Anal lobe dark maroon, capped
thinly basad by a white line.

Length of fore wing. Male, 13 mm.

clench: notes on lycaenid butterflies 215

Holotype. Male, Victoria, Mexico, February 7, 1942 (Mrs. Mary
Alice Turner), no. 26569 in the Museum of Comparative Zoology.

Remarks. From the remaining species of the group now under con-
sideration this species differs most remarkably in the utter absence of
the brilliant morpho-blue that characterizes the males. Other dis-
tinguishing characters are: the C112 tail, present only in this species
and in longula; and the spread of fulvous from the anal lobe above.
The other differences are of less importance, but can be noted by
referring to the formal description above.

This species is a striking parallel to Thecla fusius Godman and
Salvin1, which belongs to the "tailed section" of Draudt's amyntor-
group. Fusius (of which also only the male is known) is uniformly
brown above, but below is described and figured as being exactly simi-
lar to hcrodotus Fabr.2 From Godman and Salvin's figures the follow-
ing presumably significant differences have been noted : The hind wing
is longer, more produced anally (a character of the acaste group, as
opposed to the remainder ("tailed section") of the am yutor -group);
there is a very plain anal suffusion of fulvous in marialis, of which no
indication is given either in Godman and Salvin's description, or their
figure; nor is there, in marialis any indication of the bluish suffusion in
the base, as depicted in their illustration ; below there is no white line
on the hind wing of marialis, while the figure of fusius plainly shows
one, and in which fusius also agrees with hcrodotus.

There seems little doubt, therefore, that in spite of the superficial
resemblance, we are dealing with a full species, belonging even to a
separate subgroup.

The collector of this subspecies, Mrs. Mary Alice Turner, said that
she could not recall taking the specimen itself, but remembered that
collecting at Victoria wras done by an irrigation ditch, on low weeds
in open country away from the forest.

The author wishes to thank Mr. Don B. Stallings, of Caldwell,
Kansas, in whose collection the specimen formerly rested, for his kind
donation of it to the Museum of Comparative Zoology.

1 1877, Biol. Centr. Am. Lep. Rhop., 2, p. 34, pi. 52, fig. 6, 7.

2 Godman and Salvin state that the pattern below of fusius is so similar to herodotus that,
in the absence of females, they were almost inclined to regard it as a dimorphic male of that
species.

Bulletin of the Museum of Comparative Zoology
AT HARVARD COL LEG E

Vol. XCIV, No. 7

THE SOCIAL VESPIDAE OF THE GUI AN AS, PARTICU-
LARLY OF BRITISH GUIANA

By Joseph C. Bequaert

CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM

August, 1944

PUBLICATIONS
OF THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
investigations by the Staff of the Museum or of reports by spec-
ialists upon the Museum collections or explorations.

Of the Bulletin, Vols. I to XCIII, and Vol. CXIV, No. 1, 2,
3, 4, 5 and 6 have appeared and of the Memoirs, Vol. I to LVL

After 1941 no more Memoirs are to be published.

Bulletin of the Museum of Comparative Zoology

AT HA R YARD CO L L E G E

Vol. XCIV, No. 7

THE SOCIAL VESPIDAE OF THE GUIANAS, PARTICU-
LARLY OF BRITISH GUIANA

By Joseph C. Bequaert

CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM

August, 1944

Xo. 7 — The Social Vespidae of the Guianas, "particularly
of British Guiana

(Hymcnoptera)
By Joseph C. Bequaert

The present comprehensive account of the social wasps (Vespidae)
includes keys for the identification of the South American genera and
of the species known from any of the three Guianas. This area, based
on political boundaries, may be regarded as a fairly natural unit,
although the fauna is similar to that of the adjoining forests of northern
Brazil and eastern Venezuela. Its wasp fauna is, moreover, rather uni-
form and I have no doubt that all the species included in this paper
will eventually be found in British Guiana.

For the present, only the forms definitely known from British Guiana
are fully discussed, all the recorded localities of these being listed.
Social wasps live in colonies of several or many individuals and are
therefore abundant enough to be collected casually. Their nests are
also an added attraction to the general collector. Hence they are much
better known than most other Hymenoptera, with the possible excep-
tion of the social bees. Most probably few South American species
remain to be discovered, and this is particularly true of the Guianas,
one of the parts of tropical America most thoroughly investigated by
entomologists.

The Guianas are unusually rich in social wasps and would be ideal
for the study of their fascinating habits. I list from the entire area
98 structural species (and 32 additional color varieties), belonging to
16 genera (all those known from South America, except three: Synoe-
coides, Clypearia and Protonectarina). Of these, 84 species (and 18
varieties) are definitely recorded from British Guiana, 25 of them (and
8 varieties) having not yet been taken in either Dutch or French
Guiana. The wasp fauna of the British portion thus appears to be
somewhat better known than that of the others, the differences being
entirely due to insufficient collecting, in my opinion.

The following 43 species and varieties were taken at Kartabo within
the quarter square-mile area of jungle extensively studied by the New
York Zoological Society's Department of Tropical Research.

Polistes versicolor var. vulgaris
Polistes testaceicolor
Polistes melanosoma
Polistes deceptor
Mischocyttarus carbonarius

250 bulletin: museum of comparative zoology

Mischocyttarus collarellus

Mischocyttarus duckei

Mischocyttarus flavicans

Mischocyttarus labiatus

Mischocyttarus prominulus

Mischocyttarus smithii

Mischocyttarus superus

Polybia liliacea

Polybia striata

Polybia jurinei

Polybia bifasciata

Polybia bifasciata var. quadricincta

Polybia chrysothorax

Polybia micans

Polybia rejecta

Polybia occidentalis

Polybia occidentalis var. parvula

Polybia bistriata

Polybia tinctipennis var. nebulosa

Stelopolybia cajennensis

Stelopolybia pollens

Stelopolybia obidcnsis

Stelopolybia constructrix

Stelopolybia testacea

Stelopolybia pallipes var. anceps

Metapolybia cingulata

Protopolybia minutissima

Protopolybia mimdissima var. binominata

Protopolybia pumila

Apoica pallida

Apoica pallida var. arborea

Apoica pallida var. pollens

Apoica pallida var. albimacula

Brachygastra scutellaris

Synoeca surinama

Synoeca virginea

Parachartergus smithii

Sixteen additional forms listed below were taken in nearby localities,
within a fifteen-mile radius from Bartiea :

Pseudochartergus chartergoides
Mischocyttarus foveatus
Mischocyttarus heliconius
Mischocyttarus lemoulti

bequaert: social vespidae of ghe guianas 251

Mischocyttarus oecothrix

Mischocyttarus rohindicollis

Polybia catillifex

Polybia dimidiata

Polybia gorytoides

Polybia rufitarsis

Polybia sericea

Polybia singularis

Stelopolybia paraensis

Stelopolybia angulata

Stelopolybia fulvofasciata

Prot/)polybia holoxantha

Parachartcrgus fulgidipennis var. griseus

The combined two lists of 59 forms give a fair idea of the wasps to
be found in that particular section of British Guiana. Additions may
be expected, some of the more common species of the coastal lowlands,
such as Polistes canadensis and Brachygastra lecheguama, not being
included. Nevertheless, it is fairly certain that several of the Guiana
species do not occur in the Bartica area, particularly those that have
been reported only from the hilly or mountainous areas near the
Amazon watershed (Alt. Roraima, etc.).

Acknowledgments. The present study was based upon material
derived from many sources. Foremost were the collections made over
a number of years in the Kartabo district by Dr. W. Beebe and his
associates at the Tropical Research Station of the New York Zoological
Society. Additional specimens were collected by B. E. Dahlgren,
C. Geijskes, A. Mackie, the late J. G. Myers, J. Ogilvie, O. W. Rich-
ards, Neal Weber, the late W. M. Wheeler, and F. X. Williams. I
have also included Guiana records from the collections of the American
Museum of Natural History, Carnegie Museum, Cornell University
(Dept. of Entomology), Field Museum of Natural History, Museum of
Comparative Zoology and United States National Museum.

References to the original descriptions of most of the species and
color forms, and their synonyms, mentioned in this paper will be
found in Ducke's Catalogue of the Social Wasps of Brazil (1918, Rev.
Mus. Paulista, 10, pp. 313-374).

252 bulletin: museum of comparative zoology

Table of Neotropical Genera of Social Vespidae
(Based Mainly on Females and ^Yorkers)

1. Third and fourth segments of mid and hind tarsi asymmetrical,

with inner apical lobe much longer than outer one. First ab-
dominal segment much narrowed, stalk-like. . . . Mischocyttarus
All segments of mid and hind tarsi symmetrical 2

2. Thoracoabdominal muscle inserted in a narrow, slit-like furrow

at the lower end of the propodeum. First abdominal segment

short or slightly elongate, but not stalk-like Polistes

Thoraco-abdominal muscle inserted in a broadly ovate furrow at
the lower end of the propodeum 3

3. Ocelli very large, nearly as wide as basal diameter of flagellum.

Mesepisternum completely divided into an upper and a lower
plate. Abdomen long and slender, but first segment not stalk-
like Apoica

Ocelli normal or slightly swollen, much less than basal diameter of
flagellum. In doubtful cases, mesepisternum undivided 4

4. Scutellum very prominent, its vertical hind face forming an angle

with the horizontal anterior portion, which projects beyond the
short, vertical postscutellum. First abdominal segment short

and narrowed, but not stalk-like Brachygastra

Scutellum and postscutellum placed one behind the other in one
slope or convexity; scutellum sometimes slightly more raised,
but not projecting beyond postscutellum 5

5. Hind margin of postscutellum considerably extended as a sharp

angle or broad lobe into the median concavity of the propo-
deum 6

Hind margin of postscutellum either straight or forming a slight
obtuse angle or a broad curve 7

6. Clypeus usually much longer than wide, with nearly straight api-

cal margin. Outer orbits very narrow. Postscutellum divided
into a short, horizontal basal area and a vertically abrupt

apical portion Pseudochartergus

Clypeus either wider than long or at most about as long as wide,
more or less produced medially at apex. Outer orbits of normal

width. Postscutellum forming a single oblique convexity

Protopolybia

7. Labial palpi bearing a heavy, erect, curved seta some distance

from the tip. Clypeus not longer than wide, more or less pro-
duced medially at apex 8

bequaert: social vespidae of the guianas 253

Labial palpi always of 4 segments, without erect, heavy seta
before the tip. Maxillary palpi of 6 segments 10

8. Maxillary palpi of 5 segments; labial palpi of 3 segments. Small

species, with stalk-like first abdominal segment Lcipomeles

Maxillary palpi of 6 segments. First abdominal segment sometimes
narrower than second, but rarely stalk-like 9

9. Labial palpi of 4 segments Pseudopolybia

Labial palpi of 3 segments Parachartergus

10. First abdominal tergite distinctly divided into a slender basal

stalk and a broader apical portion which is part of the remaining
swollen abdomen. Outer orbits narrow. Clypeus longer than

wide, with nearly straight apical margin Charterginus

First abdominal tergite either without basal stalk or, if stalked,
the remainder of the tergite is also much narrower than the
succeeding segments and not part of them 11

1 1 . Clypeus much longer than wide, with nearly straight apical margin.

Outer orbits very narrow 12

Clypeus either wider than long or at most as wide as long, rarely
slightly longer (in some males longer than wide) ; apical margin
more or less produced, sometimes minutely bidentate. Outer
orbits normal 13

12. Abdomen subsessile, the first segment not at all stalk-like. Dor-

sum of thorax flattened. Postscutellum oblique throughout ....

Synoecoides

Abdomen with the first segment distinctly stalk-like. Dorsum of

thorax not flattened. Postscutellum with a narrow basal area,

followed by an abruptly vertical portion Clypearia

13. Abdomen subsessile, the first segment not stalk-like but posteri-

orly part of the remaining swollen abdomen. Postscutellum
with a narrow basal area slightly tuberculate in the middle,

followed by an abruptly vertical portion Chartergvs

First abdominal segment more or less stalk-like, always much
narrower than the succeeding swollen segments. Postscutellum
not so divided 14

14. First abdominal segment very long and slender, nearly as long as

thorax, with prominent spiracular tubercles Metapolybia

First abdominal segment much shorter than thorax; spiracular
tubercles weak 15

15. First abdominal segment forming a linear stalk with subparallel

sides, flattened above; second segment abruptly widened.
Clypeus with two minute apical teeth Epipona

254 bulletin: museum of comparative zoology

First abdominal segment always much wider at apex than at base.
Clypeus ending in one tooth or obtusely rounded off at apex. . 16

16. First abdominal segment with slightly prominent spiracular

tubercles ; remaining segments abruptly widened at base, sharply

conical and somewhat compressed apically Synoeca

Abdomen elongate-oval and more or less depressed 17

17. Clypeus very wide, almost twice as wide as long in the female.

Ocelli far apart, as far from one another as from the eyes

Protonectarina

Clypeus usually as wide as long or slightly longer than wide or

moderately wider; very rarely almost twice as wide as long, in

which case the ocelli are close together 18

18. Mesepisternum undivided Polybia

Mesepisternum divided by an oblique suture into an upper and a

lower plate Stelopolybia

The following names proposed for supra -specific groups of Neotropi-
cal social wasps are not given generic status in this paper.

Agelaia Lepeletier (1836), based upon Agelaia fuscicomis Lepele-
tier (1836), is at present unrecognized. The species was described
without locality. I suggest that it may have been a Polistes.

Caba R. v. Ihering (1904) = Brachygastra.

Chartergcllus J. Bequaert (1938). Subgenus of Parachartergus.

Clypeo polybia Brethes (1923) was based on a species of Mischocyt-
tarus.

Coloboclypeus Brethes (1926) is at present unrecognized. Perhaps it
was based upon a solitary vespid.

Eupolybia Dalla Torre "(1904) = Polybia.

Gymno polybia Ducke (1914) = Stelopolybia. See the discussion under
that genus.

Hypochartergus Zavattari (1906). Subgenus of Charter ginus.

Megacanthopus Ducke (1904). Subgenus of Mischocyttarus.

Melissaia Shuckard (1841) = Brachygastra.

Monacanthocnemis Ducke (1905). Subgenus of Mischocyttarus.

Myrapetra White (1841) = Polybia.

Nectarina Swainson and Shuckard (1940) = Brachygastra.

Nectarinella J. Bequaert (1938). Subgenus of Parachartergus.

Tatua H. de Saussure (1854) = E pi pona.

Xanthocaba "Cameron" Meade Waldo (1914) = Pseudo polybia.

bequaert: social vespidae of the guianas 255

Subfamily POLISTINAE

Polistes Latreille (1802)

This genus is represented in South America by more structural
species than in any other part of the World. Those known from the
Guianas or likely to occur there may be separated by the following key.

1 . Mesopleura with not even a trace of prepectal suture 2

Mesopleura at least with traces of prepectal suture (in most cases

strongly marked) 7

2. Body thickset. Abdomen ovate-fusiform, widest before mid-

length, more or less depressed apically; first tergite strongly
convex and, in profile, abruptly sloping toward the base, seen

from above as wide at apex as long or wider 3

Body slender. Abdomen elongate-fusiform, widest about mid-
length, more or less compressed apically ; first tergite moderately
convex and, in profile, gradually sloping toward the base, seen
from above longer than wide at apex 4

3. Female: head much swollen; oculo-malar space longer than one-

third of height of eye seen in front; clypeus not or barely touch-
ing eyes. Male: clypeus pentagonal, with bluntly pointed apex,
little or scarcely separated from lower inner orbits . . P. carnifex
Female: Head moderately swollen; oculo-malar space at most as
long as one-third of height of eye, usually shorter; clypeus touch-
ing inner orbits for a distance equalling one-third to one-half of
oculo-malar space. Male: clypeus subquadrate, with straight
or weakly curved anterior margin, widely separated from inner
orbits P. major

4. Propodeum very distinctly or coarsely striate 5

Propodeum finely or obsoletely striate 6

5. Occipital carina lacking over lower third of outer orbit, where the

cheek is completely rounded off into the gula; upper part of
outer orbit with an irregular slight depression near marginal

carina P. canadensis

Occipital carina continued over lower third of outer orbit, though
much weaker than in upper part, the cheek separated from the
gula by a distinct ridge; upper part of outer orbit without de-
pression near marginal carina P. apicalis

6. Occipital carina strong, continuing along lower outer orbit to base

of mandible. Humeral collar of pronotum strongly raised

P. goeldii

256 bulletin: museum of comparative zoology

Occipital carina low, absent along lower fourth of outer orbit,
where the cheek is rounded off into the gula. Humeral collar of
pronotum moderately raised P. versicolor

7. Propodeum coarsely striate. Occipital carina absent along lower

half of outer orbit, where the cheek is rounded off into the gula.
Clypeus touching eyes over a short distance in female, widely
separated from them in male. Mesepisternum divided by an

oblique suture into an upper and a lower plate P. major

Propodeum finely or obsoletely striate. Occipital carina usually
continuing along outer orbit to base of mandible 8

8. Outer orbit much widened in upper third, where the occipital

carina is strongly raised and wing-like. Male: clypeus longer
than wide, broadly rounded off at apex; all segments of flagellum

longer than wide P. occipitalis

Occipital carina not strongly raised nor wing-like at upper third of
outer orbit 9

9. Head much swollen ; outer orbit markedly wider than eye in profile ;

occipital carina slightly sinuate about mid-height. Mesepister-
num without oblique suture dividing it into an upper and a lower

plate 10

Head not swollen; outer orbit at most as wide as eye in profile;
if wider, mesepisternum partly divided by an oblique suture. 11

10. Clypeus touching eyes over a distance equal to about one-third

of length of oculo-malar space. (Head and thorax black; abdo-
men red) P. bicolor

Clypeus touching eyes over a distance equal to a little over half the
length of oculo-malar space. (Entirely oily black) . . P. deceptor

11. Body thickset. Abdomen ovate-fusiform, widest before mid-

length, more or less depressed apically; first tergite strongly
convex and, in profile, abruptly sloping toward base, seen from
above as wide at apex as long or wider. Occipital carina usually

weak along lower outer orbit near base of mandible 12

Body slender. Abdomen elongate-fusiform, widest about mid-
length, more or less compressed apically; first tergite moder-
ately convex and, in profile, gradually sloping toward base, seen
from above longer than wide at apex. Occipital carina usually
high and ridge-like along outer orbit to near base of mandible . 13

12. Mesepisternum with at least a trace of oblique suture. Clypeus of

both sexes touching eyes over a distance equal to at most one-
third of length of oculo-malar space ; lower subocular portion in
female longer than upper interocular part. Male : clypeus ending

bequaert: social vespidae of the guianas 257

in a bluntly pointed apex; thirteenth segment of antenna slightly

curved P. ruficornis

Mesepisternum without even a trace of oblique suture. Clypeus
of both sexes touching eyes over a distance equal to one-half or
more of oculo-malar space; lower subocular portion in female
about as long as upper interocular part. Male: projecting apex
of clypeus broadly rounded off; thirteenth segment of antenna
straight P. pacificus

13. Occipital carina very faint or obsolete over lower third of outer

orbit. Face of male much lengthened; clypeus higher than wide,
touching eyes over a distance equal to about one-third of length

of oculo-malar space. Thorax densely silky P. svbsericeus

Occipital carina sharp as far down as base of mandible. Face of
male not conspicuously lengthened; clypeus about as wride as
high. Thorax not densely silky 14

14. Male: Occipital carina very high and collar-like along outer orbit;

outer orbit narrower than eye in profile ; clypeus about as wide
as long; mesepisternum with a complete but very weak oblique
suture. (Head and thorax black; abdomen red. Female not

seen) P. erythrogaster

Occipital carina low, not collar-like along outer orbit 15

15. Outer orbit wider than eye in profile. Clypeus of both sexes touch-

ing eyes over a distance less than length of oculo-malar space.
Oblique suture of mesepisternum distinct in lower half, obsolete
in upper half. (Head, thorax and two basal segments of abdo-
men mostly russet with yellow markings, remainder of abdomen

black) P. testaceicolor

Outer orbit as wide as or slightly narrower than eye in profile.
Clypeus of both sexes touching eyes over a distance about equal
to length of oculo-malar space. Oblique suture of mesepisternum

lacking or very faintly indicated. (Almost entirely black)

P. melanosoma

Polistes bicolor Lepeletier (1836) is known from French Guiana,
Dutch Guiana, Brazil (Amazon Basin), Colombia and Peru.

Polistes pacificus Fabricius (1804). No doubt some of the color
forms of this species occur in British Guiana. The typical form is
known from French Guiana, as well as from Mexico, Colombia, Trini-
dad, Brazil and Peru; var. liliaciosus H. de Saussure (1854), from
French Guiana, Brazil and Peru; var. actaeon Haliday (1836) is defin-
itely known from Brazil and Paraguay, with one rather doubtful

258 bulletin: museum of comparative zoology

record from French Guiana; and var. geminatus Fox (1898) I have
seen from French Guiana and Brazil.

Polistes apicalis H. de Saussure (1858) was described from "Guiana".
There is as yet no definite record from British Guiana. I have seen it
from Guatemala, Honduras and Ecuador.

Polistes carnifex (Fabricius, 1775). This species was discussed in a
paper published in 1936 (Rev. de Entomologia, 6, pp. 376-383) It
occurs from central Mexico to northern Argentina (Misiones) in a
number of color forms, four of which I have distinguished by name.
As it is often confused with Polistes major (Palisot de Beauvois), I
have included that species also in my key, although it does not seem
to occur in the Guianas. Typical carnifex has not been reliably reported
from the Guianas, but Polistes variegata Lepeletier (1836), described
from Cayenne, may have been this form. Var. rufipennis Latreille
(1817) is known only from Panama, French Guiana (Polistes chloro-
stoma Lepeletier, 1836, from Cayenne, is a synonym) and Dutch
Guiana; and var. ochreata Spinola (1851), from Dutch Guiana, Trini-
dad, Tobago, Colombia, and Bonacca (off the coast of Honduras).
Both these color forms no doubt occur in British Guiana.

Polistes versicolor (Olivier, 1791)

One of the most common social wasps of South America, where it
varies extraordinarily in color. In a revision published in 1934 (Rev.
de Entomologia, 4, pp. 147-157), I recognized nine varieties by name.
I have since described two more (1940, Ent. News, 51, pp. 81-82).
Four of these are known from the Guianas.

1. Typical form. Only first and second abdominal tergites marked
with yellow. I have seen it from Brazil, northern Argentina, Paraguay,
Costa Rica, and British Guiana (Demerara). It was described from
French Guiana.

2. Var. vulgaris J. Bequaert (1934). The most common form of the
species, with yellow markings on at least tergites 1 to 3, often 1 to 6.
I have seen it from British Guiana (Kartabo), Panama, Colombia,
Venezuela, Trinidad, Dutch Guiana, Brazil, Paraguay, northern Ar-
gentina, Peru, Bolivia, and Ecuador.

3. Var. myops (Fabricius, 1798). Only the second tergite is marked
with yellow spots. It was originally described from French Guiana.
I have seen it from Peru. It has been recorded also from Brazil and
(perhaps by error) from Trinidad and Paraguay.

4. Var. kaieteurensis J. Bequaert (1934). Abdomen without yellow

bequaert: social vespidae of the guianas 259

markings, but the apical segments somewhat orange. Thorax with
many yellow markings. Described from British Guiana: Kaieteur.

Polistes subsericeus H. de Saussure (1854)

I have seen a specimen from British Guiana: Mt. Roraima. The
species is also known from Dutch Guiana, Brazil and Paraguay.

Polistes erythrogaster Ducke (1905)

I have seen one specimen from British Guiana: Mt. Roraima. The
species is also known from the Amazon Basin in Brazil.

Polistes testaceicolor J. Bequaert (1937)

Synonym: Vcspa analis Fabrieius, 1798 (not of Fabricius, 1775).
"Apparently a common wasp in British Guiana: Bartica; George-
town; Warina, N. W. District; Tumatumari, Potaro River; Kaieteur;
Kartabo; Mt. Everard; Arakaka, I have also seen it from Dutch
Guiana, French Guiana, Brazil (Amazon Basin), Bolivia, Colombia,
Venezuela, Peru and Costa Rica.

This species is often heavily stylopized. One female bears seven
empty pupae of Strepsiptera, protruding from the hind margins of
second (1), third (2) and fourth (2) tergites, and of third (1) and
fourth (1) sternites. One female from Bartica is labelled as taken on
carrion of agouti.

Polistes ruficornis H. de Saussure (1853)

The typical form of this species is known only from Uruguay, Para-
guay and northern Argentina. I described the var. demeraraensis
3. Bequaert (1937) from British Guiana (Mahaica River, Demerara;
Georgetown; Blairmont), and I now also refer to this form the speci-
mens from Brazil and Bolivia which I had called var. biglumoides in
1937 (Arch. Inst. Biol. Veg., Rio de Janeiro, 3, p. 180).

The unrecognized Polistes guyanensis Cameron (1912), described
from Potaro River, British Guiana, was most probably a color form of
P. ruficornis and perhaps merely a variant of var. demeraraensis.

Polistes occipitalis Ducke (1904)

British Guiana: Mt. Roraima. Also known from French Guiana,
Dutch Guiana, Brazil, Bolivia, eastern Peru, and Colombia.

260 bulletin: museum of comparative zoology

Polistes melanosoma H. de Saussure (1853)

British Guiana: Bartica; Kartabo; west bank of Demerara River.
Also known from Brazil and Paraguay.

Polistes deceptor W. A. Schulz (1905)

British Guiana: Kartabo. Also known from Dutch Guiana, Brazil
and Peru.

Polistes canadensis (Linnaeus, 1758)

This, the most widely distributed of the American Polistes, extends
from south of the Great Lakes in the United States to northern Pata-
gonia, and occurs also in some of the Lesser Antilles, but not in the
Greater Antilles. In a recent revision (1943, in process of publication),
I recognize 19 color forms; but 2 only are known thus far from the
Guianas.

1. Typical canadensis. Fairly uniformly light russet to dark
mahogany-brown; head and thorax sometimes slightly lighter than
abdomen. No yellow markings, or sometimes a narrow apical margin
on the first tergite. Wings either uniformly purplish-black or dark
russet, or more or less russet toward the tips and darker basally. This
typical form occurs in British Guiana: Demerara; Georgetown; Mt.
Roraima. It is found also in French Guiana and Dutch Guiana and
ranges from southern Arizona to northern Argentina and Rio Grande
do Sul. It does not occur in Canada.

2. Var. infuscatus Lepeletier (1836). P. canadensis amazonicus
W. A. Schulz (1905) is a synonym. Like typical canadensis, but occiput
and outer orbits more or less extensively yellow. Wings either uni-
formly purplish-black or slightly to extensively russet apically and
darker at the base. First tergite with or without narrow apical yellow
fascia. This form is perhaps more common in British Guiana than
typical canadensis: Demerara; Onverwagt. I have also seen it from
Brazil (Amazon Basin), Dutch Guiana, French Guiana, Colombia,
Panama, Peru and Ecuador.

Polistes urceolata "Klug" Erichson (1848, in Schromburgk, Reisen
in Britisch Guiana, 3, p. 590; no sex; British Guiana) was evidently
based upon some form of P. canadensis; but the description is too brief
for recognition.

bequaert: social vespidae of the guianas 261

Subfamily POLYBIINAE

Mischocyttarus H. de Saussure (1853)

Mischocyttarus is the largest Neotropical genus of social wasps,
some 73 structural species and several color forms being described to
date (1943). This number will be more than doubled in the near
future, as both Mr. O. \V. Richards and Mr. J. F. Zikan are revising
the genus and intend to describe many new forms. Meanwhile it is
possible only to enumerate those known from British Guiana.

Mischocyttarus carbonarius H. de Saussure (1853)

Megacanthopus ruficornis Cameron (1912) is probably a synonym.
British Guiana: West bank of Demerara River; Kaieteur; Kartabo.
Also known from French Guiana, Brazil, Bolivia, Peru and Panama.

Mischocyttarus cerberus Ducke (1918)

The typical form is known from Dutch Guiana and Brazil. The var.
acheron Richards (1940) was described from British Guiana: Mazaruni
Settlement.

Mischocyttarus collarellus Richards (1940)

Common in British Guiana: Mazaruni Settlement; Cuyuni River, 3
miles from Kartabo; Potaro River, on trail between Tukeit and Kaie-
teur; Moraballi Creek, Essequibo River; Kartabo; Aremu, Bartica
District. Also known from Dutch Guiana, Brazil and Panama.

Mischocyttarus collaris Ducke (1904)

British Guiana: Berbice Savannas; Courantyne River; Demerara.
Also known from Brazil.

Mischocyttarus duckei R. du Buysson (1909)
British Guiana : Kartabo. Originally described from French Guiana.

Mischocyttarus flavicans Fabricius (1804)

Megacanthopus goeldii Ducke (1905) is a synonym.
British Guiana: Kartabo; Mazaruni River. Also known from
French Guiana, Dutch Guiana, Brazil, Bolivia, Peru and Ecuador.

262 bulletin: museum of comparative zoology

Mischocyttarus foveatus Richards (1940)

Described from British Guiana: First Falls of Essequibo River;
Issororo, Northwest District; Island near Monkey Jump; Moraballi
Creek, Essequibo River.

Mischocyttarus heliconius Richards (1941)

Described from British Guiana: Moraballi Creek, Essequibo River;
Potaro River, on trail between Tukeit and Kaieteur.

Mischocyttarus injucundus (H. de Saussure, 1854)

The typical form is known from Brazil and Colombia. The var.
bimarginatus (Cameron, 1912) was described from British Guiana,
without more definite locality. I have seen it from Demerara; Maza-
runi River; Blairmont; and Georgetown

Mischocyttarus labiatus (Fabricius, 1804)

Megacanthopus atriceps Cameron (1912) is a synonym.

British Guiana: Mt. Roraima; Rupununi River; Kaieteur; west
bank of Demerara River; Kartabo; Bartica. Also known from French
Guiana, Dutch Guiana, Trinidad, Brazil, Venezuela, Paraguay, Peru,
Colombia and Panama.

Mischocyttarus lecointei (Ducke, 1904)

British Guiana : Tukeit ; Kaieteur. Also known from French Guiana,
Dutch Guiana and Brazil.

Mischocyttarus lemoulti (R. du Buysson, 1909)

British Guiana: Moraballi Creek, Essequibo River. Originally de-
scribed from French Guiana.

Mischocyttarus metathoracicus (H. de Saussure, 1854)

British Guiana: Essequibo River; Oko River, a tributary of the
Cuyuni. Originally described from French Guiana; also known from
Brazil and Peru.

bequaert: social vespidae of the guianas 263

Mischocyttarus metoecus Richards (1940)

Described from British Guiana: Mazaruni Settlement. Also known
from Dutch Guiana and Brazil.

Mischocyttarus oecothrix Richards (1940)

Described from British Guiana: Kaieteur Savanna, Potaro Valley;
Canister Falls, Cattle Trail Survey; Moraballi Creek, Essequibo River.

Mischocyttarus prominulus Richards (1941)

Described from British Guiana: Kartabo; Mazaruni Settlement.
Also known from Bolivia.

Mischocyttarus rotundicollis (Cameron, 1912)

Originally described from British Guiana, without more definite
locality. I have seen it from Kalacoon and Penal Settlement, Bartica
District.

Mischocyttarus smithii H. de Saussure (1853)

Megacantkopus violaceipennis Cameron (1912) is a synonym.
British Guiana : Kartabo ; Essequibo River. Also known from French
Guiana and Brazil.

Mischocyttarus superus Richards (1940)

Described from British Guiana: Mazaruni Settlement; Bartica-
Potaro Road; Potaro River, on trail between Tukeit and Kaieteur;
I have also seen it from Kartabo.

Mischocyttarus surinamensis (H. de Saussure, 1854)

British Guiana: Kamakusa; Courantyne River; Trail between
Tukeit and Kaieteur; Demerara. Also known from Dutch Guiana and
Brazil.

Mischocyttarus synoecus Richards (1940)

Described from British Guiana: Amatuk, Potaro River; Mazaruni
Settlement. Also known from Brazil and Panama.

Polybia nigriceps Cameron (1912), described from British Guiana,
without more definite locality, was a species of Mischocyttarus. The

264 bulletin: museum of comparative zoology

name is preoccupied by Polybia fastidiosuscula var. nigriceps Zavattari
(1906).

Mr. O. W. Richards (in litt.) includes British Guiana in the range of
Mischocyttarus drewseni (de Saussure, 1857).

Mischoeyttarus socialis (H. de Saussure, 1854) (= Vespa atra Olivier,
1791; not of Gmelin, 1790) was taken in French Guiana, Cayenne
being the type locality of Olivier's V. atra.

Mischocyttarus alfkenii (Ducke, 1904) I have seen from Dutch
Guiana.

Pseudochartergus Ducke (1905)

I have revised this genus in a recent paper (1938, Rev. de Entomo"
logia, 9, pp. 103-105), where I recognized only two species. A study
of more extensive material in the collections of Cornell University has
led me to separate a third species, Pseudochartergus panamensis (Zavat-
tari, 1906) ( = Chartergus acutiscutis Cameron, 1907). This differs from
P. chartergoides in the relatively shorter clypeus of the female (about
as wide as long) and in the lateral ridges of the propodeum being more
rounded off. All localities from British Honduras, the Republic of Hon-
duras, Panama and Colombia, which I listed in 1938 under P. charter-
goides var. cinctellus, refer to P. panamensis.

Only one of the three species is known from the Guianas.

Pseudochartergus chartergoides (Gribodo, 1891)

I distinguish by name three color forms of this species, but only the
var. cinctellus (Fox, 1898) occurs in British Guiana: Kartabo. It is
known also from Dutch Guiana and Brazil.

Charter ginus pallidibalteatus Cameron (1912), described from
British Guiana, was most probably P. chartergoides var. cinctellus.

Charterginus Fox (1898)

This genus was revised in 1938 (Rev. de Entomologia, 9, pp. 99-
103). Only one of the four species is known from the Guianas.

Charterginus huberi Ducke (1904)

British Guiana: Source of Essequibo River; Waratuk. It was de-
scribed from the Amazon Basin of Brazil. I have seen it also from
Dutch Guiana and French Guiana (Maroni). Polybia fuhicauda

bequaert: social vespidae of the guianas 265

Cameron (1912), described from British Guiana, is a synonym of
C. huberi.

Protopolybia Ducke (1905)

This genus, containing some of the smallest South American wasps,
is well represented in the Guianas. In addition to the species included
in the key, three forms described by Cameron from British Guiana
are as yet unrecognized.

Key to Guiana Species

1. Concavity of propodeum narrow, groove-like. First abdominal

tergite slender, the basal third or more stalk-like. Clypeus about

as wide as high 2

Concavity of propodeum broad, either very shallow or deeply
bowl-shaped. First abdominal tergite thickset, not or very briefly
stalk-like at base 3

2. Thorax rather dull, with distinct, scattered, medium-sized punc-

tures; abdomen impunctate. Postscutellum nearly twice as wide
as greatest length, with triangular, bluntly pointed apical exten-
sion. (Extensively and fairly uniformly testaceous-yellow ; almost

bare) P. holoxantka

Body dull or slightly shiny, usually with a few discrete larger
punctures. Postscutellum at most one and one-half times as wide
as greatest length, with the apical extension rounded off, tongue-
like. (Black, with few or many yellowish markings)

P. minutissima

3. First abdominal tergite very broad and short, cap-shaped, passing

gradually into the base of the second, which is not constricted
from it. Outer orbit unusually widened in lower half; occipital
carina low and ending some distance from base of mandible.
Clypeus nearly twice as wide as high. Thorax thickset, about as
high as long, with many coarse punctures; humeral collar high,
shouldered at the sides; postscutellum nearly twice as wide as
long; sides of propodeum bulging, somewhat compressed, though

broadly rounded off P. emortualis

First abdominal tergite not cap-shaped, always set off from the
more swollen second. Thorax longer than high; sides of propo-
deum low, depressed, though broadly rounded off 4

4. Clypeus one and one-half times to nearly twice as wide as high,

smooth and shiny. Outer orbit about as wide as eye, not reced-

266 bulletin: museum of comparative zoology

ing; occipital carina high, extending to near base of mandible.
Concavity of propodeum deep; postscutellum about one and one-
half times as wide as long; humeral collar high, more or less
shouldered at the sides. Body very shiny, with distinct, scattered,
medium-sized to large punctures, with many long, erect hairs

P. picteti

Clypeus much less than one and one-half times as wide as high.
Outer orbit narrower than eye, receding. Postscutellum slightly
though distinctly shorter than wide. Body dull or nearly dull;

erect hairs short and sparse 5

5. Thorax about one and one-fourth times as long as high. Body dull,
fairly uniformly covered with small, discrete punctures. Clypeus
about one and one-third times as wide as high. (Mostly pale
yellow) P. amarella

Thorax nearly one and one-half times as long as high. Body not or

slightly shiny, with microscopic, alutaceous sculpture and a few

small, discrete punctures. Clypeus at most one and one-fourth

• times as wide as high. (Black, marked with yellow) . .P. pumila

Protopolybia picteti (H. de Saussure, 1854) is a widely distributed
species, of which I recognize by name 9 color forms. Only one of these
has been reported from the Guianas. P. picteti var. bella (R. von Iher-
ing, 1903) was originally described (as Polybia bella) from Dutch
Guiana. It is also known from Brazil, Peru and Bolivia. It is char-
acterized by the ground color of the body being black or slightly
brownish-black, with many yellow markings on thorax and abdomen
(usually no yellow stripes on mesonotum) and 3 pale spots on the base
of tergite 2, the median spot being transverse (wider than long).

Protopolybia holoxantha (Ducke, 1904)

This species is probably rather common in British Guiana: Kama-
kusa; Moraballi Creek, Essequibo River; Penal Settlement, Bartica
District. It is also known from French Guiana and Brazil (Oyapoc
and Amazonas).

Protopolybia minutissima (Spinola, 1851)

I recognize four color forms of this species in the Guianas.

1. Pale markings few, whitish or very pale yellow; base of tergite 2
without cross-band or lateral spots 2

bequaert: social vespidae of the guianas 267

Pale markings more extensive, pale to bright yellow; base of tergite
2 with a cross-band or a pair of spots 3

2. Thorax not or very little marked with pale yellow or whitish; no

spots on propodeum typical minutissima

Thorax more profusely marked with pale yellow or whitish; a pair
of spots on propodeum var. bitwminata

3. Body profusely marked with yellow; base of tergite 2 with a con-

tinuous and usually broad cross-band var. sedula

Body less extensively marked with yellow; base of tergite 2 with a
pair of streaks or spots var. exigua

The typical form occurs in British Guiana (Kartabo), as well as in
Peru and Brazil.

The var. binominata (YV. A. Schulz, 1906) appears to be the most
common form, being known from British Guiana (Kartabo; George-
town; Demerara), Colombia, Panama, Ecuador, Peru, and Bolivia.
Polybia minutissima H. de Saussure (1854) was this form, not Spinola's
typical minutissima.

The var. exigua (H. de Saussure, 1854) is fairly common in Brazil
and has been reported from French Guiana, Trinidad, and eastern
Peru (Iquitos).

The var. sedula (H. de Saussure, 1854) is widespread and has been
taken in Dutch Guiana, French Guiana, Brazil, Venezuela, Trinidad,
Colombia, Panama (Darien), Peru, Bolivia, and Paraguay. I regard
Polybia diligens F. Smith (1857) as a synonym of sedula.

Protopolybia amarella, new species

Fig. 1

Female. Head flattened, slightly wider than thorax; seen in front,
subcircular, scarcely wider than high; from above, rectangular with
receding hind corners, about two and one-half times as wide as long;
occipital margin with a broad and shallow inward curve. Vertex and
genae margined by a fine carina, which ends a short distance from the
base of the mandible ; gena slightly narrower than eye in profile. Oculo-
malar space lacking. Inner orbits about one and one-half times as far
apart on vertex as at clypeus. Ocelli fairly large, in a nearly equilateral
triangle ; posterior ocelli slightly farther from eyes than from each other.
Interantennal shield broad, not set off, with a longitudinal groove over
upper half; antennae somewhat farther apart than from eyes. Frons
slightly convex. Clypeus at its narrowest about one and one-third

268

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times as wide as high, irregularly heptagonal, contiguous to the eyes
over about one-half of sides; apical margins moderately produced,
ending in a blunt, rounded point. Mandible with an oblique cutting
edge of four sharp, subequal teeth. Antenna: scape long, slender,
slightly curved; second segment unusually long, only slightly shorter
than third, scarcely swollen; remainder of flagellum slightly club-

Fig. 1. Protopolybia amarella J. Bequaert. Female. A, body in profile; B,
first and second tergites from above; C, postscutellum and propodeum from
behind; D, head in front view.

shaped. Thorax rather thickset; in profile, about one and one-fourth
times as long as high; from above, nearly one and two-thirds times as
long as wide before tegulae. Pronotum nearly straightly truncate
anteriorly; humeral margin with a distinct obtuse ridge which stops at
the sides, forming slight, blunt humeral angles; dorso-lateral areas
sloping; lower anterior margin of ventro-lateral areas with a swelling
indented by a deep pronotal fovea. Mesopleuron much swollen, with-
out mesepisternal suture; deep mesepimeral suture ending abruptly

bequaert: social vespidae of the guianas 269

and far from metapleuron. Upper sclerite of metapleuron ending
nearly square below, not extended along meso-metapleural suture.
Scutellum slightly and evenly convex, with a very weak median
impressed line. Postscutellum triangular, scarcely swollen, its median
area about one and one-third times as wide as long, uniformly sloping
or nearly vertical throughout, the lower extension very long, with a
free, raised, tongue-like apex. Propodeum little swollen, nearly vertical
in profile; median concavity very broad and shallow. First abdominal
tergite short and cup-shaped, with a very short, broad stalk, which is
not wing-like at the sides; seen from above, gradually widened, slightly
shorter than wide at apex, about half as wide at apex as second seg-
ment; in profile, very gradually and moderately convex. Second
tergite from above wider than long, gradually widened at base ; in pro-
file, moderately and evenly convex. Apical margins of all segments
very slightly thickened. Mid tibiae with two spurs. Claws symmetri-
cal, unarmed. Venation: second cubital cell much higher than wide;
third cubital long, about one and one-third times as long on cubitus
as on radius ; radial cell very long, acute ; basal vein ending in subcosta
Hose to the large stigma.

Dull; fairly uniformly covered with small, discrete punctures, more
weakly on frons; clypeus and mandibles almost impunctate. Pubes-
cence sparse, short, erect, somewhat longer on propodeum and abdo-
men. Eyes bare.

Pale yellow, with ferruginous blotches on antennae, tip of mandible,
tibiae, tarsi and under side of abdomen. A few brownish-black spots
as follows : a streak behind each posterior ocellus ; a transverse row of
four spots on the prehumeral depressed slope of pronotum; narrow
sutures of mesonotum, scutellum and postscutellum; a median line
over anterior half, a line on each side over posterior half, and a pair of
elongate spots on disk of mesonotum. Disk of second tergite with a
median blackish triangle showing through the yellow integument.
Wings hyaline, with a yellowish tinge; veins and stigma pale yellowish-
testaceous.

Length (h.+th.+t. 1+2): 5.1 mm.; of fore wing, 5 mm.
British Guiana: Source of Essequibo River, female holotype (J.
Ogilvie). — Colombia: La Chorrera, Rio Igara-Parana, Intendencia
Amazonas, female paratype (J. C. Bradley). — Holotype at Mus.
Comp. Zool., Cambridge, Mass.; paratype at Dept. Ent., Cornell
University.

The almost completely yellow body makes this a striking insect.

270 bulletin: museum of comparative zoology

Protopolybia emortualis (H. de Saussure, 1855)

This species occurs in two color forms.

1. Typical emortualis is more or less extensively ferruginous, partic-
ularly on the abdomen, and has pale yellowish markings. It was de-
scribed from Brazil and Ducke reported it from Dutch Guiana.

2. The var. duckei (R. du Buysson, 1905) ( = Charter -ginus duckei
R. du Buysson) differs only in the ground color of the body, including
the abdomen, being black. It was originally described from Brazil
(Bahia) ; but Ducke reports it from eastern Peru (Iquitos) and I have
seen it from British Guiana (Upper Essequibo River). It is doubtful
whether this form is worth separating from typical emortualis.

Protopolybia pumila (H. de Saussure, 1863)

Two forms of this species are known in the.Guianas.

1. Typical pumila is extensively marked with yellow, the meso-
notum often bearing yellow stripes and the base of tergite 2 a pair of
unconnected spots. I have seen it from British Guiana: Kartabo;
source of Essequibo River. It is also known from Dutch Guiana,
Brazil, Venezuela, Colombia, Peru, and Bolivia.

2. The var. rotimdata Ducke (1910) was described (as a species)
from French Guiana and is not known from elsewhere. It has few pale
markings, no stripes on mesonotum nor spots at the base of tergite 2.

Protopolybia rufo-ornata (Cameron, 1912)

This was described, as Charter ginus rufo-ornatus, from British
Guiana, without more definite locality. It is at present unrecognized.
No doubt it was a Protopolybia, possibly a color form of P. picteti.

Protopolybia sulciscutis (Cameron, 1912)

This also was described merely from "British Guiana," as Polybia
sulciscutis, and is unrecognized. The statement in the description,
"apex of postscutellum narrowed to a broad, bluntly rounded point,"
seems to place it in Protopolybia.

Protopolybia nana (Cameron, 1912)

This is another unrecognized species described by Cameron, as
Polybia nana, from "British Guiana." The name conflicts with the

bequaert: social vespidae of the guianas 271

earlier Polybia nana H. de Saussure (1803); hut, as Cameron's wasp is
probably identical with some other described Protopolybia, it would be
premature to rename it.

Brachygastra Perty (1833)

This genus has been generally called Ncctarina Swainson and
Shuckard (1840) ; but there seems to be no reason why the earlier name
Brachygastra should not be used.

Key to Guiana Species

1. Sides of propodeum compressed but not angular, forming evenly

rounded curves in profile. Head (except clypeus), thorax and
second abdominal tergite with deep, large punctures. Dorsal
(horizontal) area of scutellum four times as wide as long in the

middle B. Scutellaria

Sides of propodeum compressed into prominent, broad, blunt angles.
Second abdominal tergite with small punctures only 2

2. Dorsal (horizontal) area of scutellum nearly flat, less *han three

times as wide as long in the middle, with the hind margin rather
sharp but slightly or scarcely curved inward. Punctures of meson-
otum fairly numerous and not or scarcely stronger than those of
the base of second tergite. Clypeus about twice as wide as

high B. lecheguana

Dorsal (horizontal) area of scutellum convex, three to four times as
wide as long in the middle, with the hind margin blunt but deeply
curved inward (scutellum forming two broad lobes). Punctures
of mesonotum decidedly stronger than those of second tergite . . 3

3. Body with many erect, short hairs. Punctures of frons and meso-

notum more numerous and fairly close. Lateral angles of pro-
podeum higher. (Usually with many yellow markings)

B. bilineolata

Body with very few erect hairs. Punctures of frons and mesonotum

far apart. Lateral angles of propodeum lower. (With few yellow

markings, the thorax and head almost entirely black)

B. augusti

Brachygastra augusti (de Saussure, 1854)

The typical form of the species has distinct yellow apical bands on
tergites 1 to 6 and sternites 2 to 6 of the abdomen. It is known from

272 bulletin: museum of comparative zoology

British Guiana (source of Essequibo River), Dutch Guiana, French
Guiana, Brazil, Paraguay, Bolivia, Peru, Colombia, and Venezuela.

Brachygastra lecheguana (Latreille, 1824)

This widely distributed and common wasp, known from southern
Arizona and southern Texas to northern Argentina, was discussed in
a paper published in Entomologica Americana in 1932 (13, pp. 92-
112). The specimens from British Guiana (Arakaka), Dutch Guiana
and French Guiana are of the var. velutina Spinola (1841), which has
the thorax densely covered with silky, golden-yellow hairs.

Brachygastra scutellaris (Fabricius, 1804)

The several color forms of this species are revised in Jr. New York
Ent. Soc, 50, (1942) 1943, pp. 306-308, where I recognize six of them
by name. The three known from the Guianas may be separated as
follows :

1. Scutellutn black; postscutellum either black or with an anterior

pale cross-band. Abdomen with whitish apical margins on most

sternites and some of the tergites var. myersi

Scutellum and postscutellum mostly or entirely yellow 2

2. Second abdominal tergite mostly orange-yellow or ferruginous.

Thorax with many yellow markings ; mesonotum often with spots

or stripes var. rufiventris

Second abdominal tergite with only an apical yellow margin. Yel-
low of pronotum either lacking or restricted to the narrow, often
incomplete humeral margin typical scutellaris

Typical scutellaris is the most common form in British Guiana:
Forest Settlement, Mazaruni River; Kartabo; junction of Mazaruni
and Essequibo Rivers. Ducke lists it from Bartica, and Bodkin from
Issororo, N. W. District. It is also known from French Guiana, Brazil,
Bolivia, eastern Peru, Colombia, and Panama. Brachygastra scutellata
Spinola (1851) is a synonym of the typical form.

The var. rufiventris (de Saussure, 1854) is reported from French
Guiana, Brazil and Colombia.

The var. myersi J. Bequaert (1943) was described from British
Guiana (Mt. Roraima) and Bolivia.

bequaert: social vespidae of the guianas 273

Brachygastra bilineolata (Spinola, 1841)

Recently (1943, Jr. New York Ent. Soc, 50, for 1942, pp. 303-306)
I recognized five color forms of this species, those occurring in the
Guianas being separated in the following key:

1. Second tergite with only the apical margin yellow. Mesonotum

with short yellow longitudinal stripes or a pair of yellow spots
posteriorly, or entirely black. Wings with a honey-yellow tinge,

most of the veins testaceous typical bilineolata

Second tergite with a broad or narrow discal yellow band in addi-
tion to the apical fascia, or mostly yellow. Mesonotum always
with two yellow stripes 2

2. Second tergite mostly yellow, except for the black base and a more

or less defined transverse black discal blotch or irregular line.

Wings with a honey-yellow tinge var. Surinam ensis

Second tergite with two separate transverse yellow stripes of about
equal width, one apical, the other discal 3

3. Wings extensively tinged with honey-yellow. Veins and stigma

mostly pale var. antillarum

Wings nearly hyaline, slightly grayish, not suffused with yellow.
Veins and stigma blackish var. smithii

Typical bilineolata is known from British Guiana (Demerara;
Georgetown), French Guiana, Dutch Guiana, Venezuela, Brazil,
Colombia, and Costa Rica. I regard Nectarinia mobiana de Saussure
(1867) as a synonym of typical B. bilineolata.

The var. antillarum (Provancher, 1883) occurs in British Guiana:
Rapununi River. I have seen it also from Dutch Guiana, Trinidad,
Brazil, eastern Peru, and the Republic of Honduras.

The var. smithii (de Saussure, 1854) has been found in French
Guiana, Dutch Guiana, Brazil, and Colombia.

The var. surinamensis J. Bequaert (1943) is known only from Dutch
Guiana.

Chartergus Lepeletier (1836)

As stated under Parachartergus, the type of this genus was desig-
nated by Emile Blanchard (1840) as Vespa nidulans Fabricius (1793),
a synonym of Vespa chartaria Olivier (1791). I revised the genus in
1938 (Rev. de Entomologia, 9, pp, 113-115) under the erroneous name
Epipona. I recognize two species, of which only one occurs in the
Guianas.

274 bulletin: museum of comparative zoology

Chartergus chartarius (Olivier, 1791)

I have seen this wasp from British Guiana: Demerara; Georgetown.
It occurs also in French Guiana, Dutch Guiana, Brazil, Paraguay,
Bolivia, and Peru (Iquitos).

Synoeca H. de Saussure (1852)

Ducke (1910) recognizes only two species of Synoeca, but I believe
three may be separated on structural characters, as du Buysson did
in his Monograph (1906). Moreover, Polistes virginea Fabricius
appears to be the valid name for the species usually called S. irina
(Spinola).

1. Larger; fore wing 16 to 18 mm. long. Eyes bare. Clypeus smooth

and shiny. Propodeum either impunctate or with a few minute
punctures. Oculo-malar space of worker and queen fully as long
as sixth antennal segment. Mostly or wholly bluish-black; wings

violaceous-black S. surinama

Smaller; fore wing 13 to 15 mm. long. Eyes with a few short hairs.
Clypeus rather dull and with scattered punctures. Propodeum
densely punctate. Oculo-malar space of worker and queen
shorter than sixth antennal segment. Either bluish-black or
partly ferruginous. Wings mostly yellowish-subhyaline 2

2. Thorax fairly uniformly punctate, the punctures of mesopleura and

mesonotum scarcely smaller than those of propodeum. Scutellum
convex, with very slight median impressed line. Swollen portion
of first abdominal segment elongate-triangular seen from above.
Black costal border sharply set off from the remainder of the

wings S. chalybea

Punctures of mesopleura and mesonotum very fine, much weaker
than those of propodeum. Scutellum gibbose, with a deep median
saddle. Swollen portion of first abdominal segment trapezoidal

seen from above. Wings fairly uniformly yellowish-russet

S. virginea

Synoeca surinama (Linnaeus, 1767)

A common wasp throughout Central and South America, from
southern Mexico to southern Brazil. The size of the head varies con-
siderably in this species and seems to be correlated with the total size
of the specimen. Perhaps these differences set off the fertile females
(queens) from the workers. Several other insects are homoeochromic

bequaert: social vespidae of the guianas 275

with S. surinama. The most interesting of these is the mantispid,
Climaciella chalybea Erichson, of which I have seen a specimen from
the Upper Rio Huallaga, Peru. The collector had evidently mistaken
it for the wasp. Possibly the mantispid lives in the nest of Synoeca.

1. Typical form. Nearly entirely bluish-black, at most with russet
blotches on the mandibles. This is Vespa .surinama Linnaeus (1767),
described from Surinam, where it is common. Vespa nigricornis
Olivier (1791), Polistcs cacrulea Fabricius (1804) and Synoeca ultra-
marina H. de Saussure (1852) are synonyms.

Common in British Guiana: Arakaka; Penal Settlement, Bartica;
Rockstone, Essequibo River; -Kartabo; Georgetown; Kamakusa;
Berbice Savannas; Blairmont. I have also seen it from Costa Rica,
Colombia, Venezuela, Trinidad, French Guiana, Dutch Guiana, Brazil,
Ecuador, Peru, and Bolivia.

2. Var. cyanea (Fabricius). The lower part of the head (particularly
the clypeus) is more or less ferruginous-red; second abdominal tergite
occasionally blotched with russet. Specimens transitional to typical
surinama are frequent. This is Vespa cyanea Fabricius (1775), Synoeca
azurea H. de Saussure (1S52) and Synoeca violacca H. de Saussure
(1852).

This form is more widely distributed than typical surinama and is
known from French Guiana. I have seen it also from Mexico (Apat-
zingan, State of Michoacan), the Republic of Honduras, Guatemala,
Costa Rica, Panama, Colombia, Brazil, Paraguay, Ecuador, and
Venezuela.

Synoeca virgixea (Fabricius, 1804)

This species is rarer and not as widely distributed as S. surinama,
being apparently restricted to the northern half of South America.
It is almost entirely ferruginous or russet with very slight or no bluish
sheen. Fabricius' description of Polistcs virginea (1804) fits this wasp
exactly. The statement "Antennae nigrae articulo secundo [ = scape]
longiore, testaceo", agrees with the Synoeca, not with Apoica pallida.
Moreover, W. A. Schulz (1912, Berlin Ent. Zeitschr., 57, p. 84) recog-
nized Synoeca irina in Fabricius' type. Polistcs irina Spinola (1851)
and Synoeca testacea H. de Saussure (1854) are synonyms.

British Guiana: Kartabo; Junction of Mazaruni and Essequibo
Rivers; Penal Settlement, Bartica District; Source of Essequibo River.
I have also seen it from Dutch Guiana, Brazil, Ecuador, and Peru.

276 bulletin: museum of comparative zoology

Synoeca chalybea H. de Saussure (1852)

1. The typical form of this species, as figured by de Saussure (1854),
is nearly entirely bluish-black, with or without violaceous reflections ;
mandibles, clypeus, oculo-malar spaces and spots on mesopleura and
legs are yellowish-russet. The specific name was originally spelled
chalibea, but later (1854) corrected to chalybea by the author himself.

British Guiana: Shudihar River. I have also seen it from Brazil and
Peru and it was originally described from French Guiana.

2. S. chalybea var. splendens R. du Buysson (1906) was originally
described as a form of S. irina; but all specimens I have seen have the
puncturation of the thorax and the shape of the first abdominal seg-
ment as in S. chalybea. It is more or less extensively ferruginous or
yellowish-russet. I have seen it from Bolivia and Peru. As Dr. Wey-
rauch collected both typical chalybea and var. splendens apparently
from the same nest at Satipo, Peru, it is doubtful whether the name
splendens deserves to be retained.

Metapolybia Ducke (1905)

I have reached the conclusion that there are two structural species
in this genus, not one as Ducke claimed. They may be separated as
follows :

1. First abdominal segment slightly widened apically seen from
above and, in profile, very gradually thickened over posterior

half. Ground color of abdomen reddish-brown M . suffusa

First abdominal segment distinctly widened apically seen from
above and, in profile, rather abruptly thickened over posterior
third. Ground color of abdomen black M. clngulata

M. suffusa (Fox, 1899) has not yet been taken in the Guianas.
According to Fox' types from Corumba, Brazil, this is the wasp with
the abdomen extensively russet which Ducke (1910) mentions from
Bolivia and calls Metapolybia pediculata var. rufopicta. I have also
seen it from Peru, Bolivia and Colombia. At Restrepo (Int. Meta),
Colombia, I found M. suffusa and clngulata nesting side by side, but in
distinct colonies. They were not in the least aggressive, although the
sting was painful.

Metapolybia cingulata (Fabricius, 1804)

Schulz (1912) saw the types of Eumenes cingulata Fabricius (1804)
at Copenhagen and recognized in them Polybia pediculata H. de

bequaert: social vespidae of the guianas 277

Saussure (1854). The extent of pale yellow markings varies, Fabricius'
cingulata being based upon specimens with only the first and second
tergites margined with yellow. This is also true of Gribodo's Tatua
decorata (1896); while Polybia pediculata var. unilineata R. v. Ihering
(1904) refers to specimens with only traces of yellow markings. There
is no name available, and none seems needed, for specimens with most
or all tergites margined.

British Guiana: Kartabo; Bartica. Also known from Venezuela,
Trinidad, Dutch Guiana, French Guiana, Brazil, Paraguay, northern
Argentina, Bolivia, Peru, Colombia, Panama, Costa Rica, Guate-
mala, and Mexico (Cordoba, Vera Cruz; Tierra Colorada, Guerrero).

Epipona Latreille (1802)

Emile Blanchard in 1840 (Hist. Nat. Ins., 3, Orth. Nevr. Hem. Hym.
Lep. Dipt., p. 394) designated as type of Epipona, Vespa morio
Fabricius (1798), a synonym of Vespa tatua Cuvier (1797). This gen-
eric name must therefore be used instead of Tatua de Saussure (1854) ;
while the genus I called Epipona in 1938 should be known as Charter-
gus Lepeletier. Epipona was revised in 1938 (Rev. de Ehtomologia, 9,
pp. 115-117), when I recognized two species, only one of which occurs
in the Guianas.

Epipona tatua (Cuvier, 1797) has not yet been taken in British
Guiana; but it is known from French Guiana, Trinidad, Venezuela,
Brazil, Peru, Colombia, Panama, and Costa Rica.

Polybia Lepeletier (1836)
Key to Guiana Species

1. Propodeum with more or less distinct transverse striation extend"

ing even over the convex sides. Occipital margin of head with a

deep inward curve. Body 14 to 16.5 mm. long 2

Propodeum without or with obsolete striation; rarely a few irregu-
lar striae in the concavity or on the sides 4

2. Head very strongly swollen behind the eyes; outer orbit slightly

wider than eye in profile. Clypeus broader than high, the apex
evenly and broadly rounded off. Pronotum on the sides with a
strong but blunt humeral crest. Striation of propodeum fine,

but regular P. ujhelyii

Head moderately swollen behind the eyes; outer orbit narrower
than eye in profile. Clypeus about as high as wide, with bluntly
pointed apex. Pronotum somewhat shouldered, but not ridged

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at the humeral angles. Striation of propodeum very irregular
or partly effaced 3

3. Concavity and sides of propodeum with coarse and irregular

wrinkles and punctures. First abdominal tergite slightly swol-
len; stalk-like base passing gradually into swollen apical por-
tion (in profile view) P- striata

Striation and punctures of propodeum weak or obsolete. First
abdominal tergite strongly swollen, abruptly raised behind the
stalk-like base (in profile view) P. liliacea

4. Median concavity of propodeum distinct and more or less circular,

never longer than wide 5

Median concavity of propodeum either almost lacking or groove-
like and longer than wide 7

5. Head distinctly swollen; outer orbit fully as wide as eye in profile.

Puncturation of mesopleura fine but distinct. Black, with
scutellum and postscutellum entirely yellow. Body 12 to 14

mm. long P • jurinei

Head not swollen; outer orbit narrower than eye in profile. Meso-
pleura nearly impunctate. Smaller; body 8 to 10 mm. long.. .6

6. Pronotum with very weak, blunt humeral collar; hind margin

forming an oval curve. First abdominal segment slender;

swollen portion longer than wide at apex P. signata

Pronotum with distinct, sharp humeral collar; hind margin nearly
semi-circular. First abdominal segment shorter; swollen portion
about as long as wide at apex P- bifasciata

7. At least sides of thorax covered with dense, silky pubescence,

often with golden sheen. Yellow markings lacking or much

reduced. Body 14 to 19 mm. long 8

Thorax nowhere covered with dense, silky pubescence 11

8. Eyes with distinct short hairs. Oculo-malar space very long (as

long as fifth antenna! segment). Pronotum without humeral
collar. Black, with thorax more or less extensively and first or
first and second abdominal segments reddish-brown. Body 15

to 17 mm. long P- sericea

Eyes bare. Oculo-malar space very short or lacking 9

9. Dorsum of thorax densely covered with long golden-silky pubes-

cence. Mesopleura finely but distinctly punctate. Pronotum
without humeral collar. Mostly brownish-red to fuscous;
abdomen without paler apical margins; head black; wings more
or less ferruginous, particularly toward costal margin. Clypeus
about as long as wide. Body 15 to 17 mm. long. . P. chrysothorax

bequaert: social vespidae of the guianas 279

Dorsum of thorax only sparsely pubescent, not golden-silky.
Color different 10

10. Only pleura of thorax distinctly silky; the abdomen not. Wings

subhyaline or slightly brownish-yellow, with ferruginous costal

margin. Body 15 to 16 mm. long P. affinis

Entire body, including abdomen, silky or velvety. Either russet
to brownish-russet with testaceous apical abdominal margins,
or (var. velutiiia) almost entirely blackish without apical mar-
gins. Wings strongly tinged with honey-yellow. Thorax very
finely punctate. Pronotum without humeral collar. Clypeus
slightly wider than long. Body 14 to 18 mm. long. . . P. micans

11. Large species, 18 to 21 mm. in total length. Head and thorax

black; abdomen red; wings nearly hyaline. Thorax shiny, with
scattered punctures and long, erect hairs. Oculo-malar space

long. Clypeus much wider than long P. dimidiata

Smaller. Thorax not shiny, or the other characters mentioned
above not all present 12

12. First tergite distinctly punctate. Mesopleura and propodeum

with large punctures. Outer orbits somewhat swollen in upper
half. Clypeus slightly wider than high. Eyes with distinct short
hairs. Body black, somewhat velvety, 12 to 14 mm. long; wings

infuscate, clearer posteriorly or apically 13

First tergite usually impunctate; if distinctly punctate, the other
characters mentioned above not all present 14

13. Oculo-malar space at its shortest over half the length of the

tenth antennal segment. Concavity of propodeum narrow,
groove-like, but very weak or barely indicated in upper third

P. tincti'pennis
Oculo-malar space at its shortest less than half the length of the
tenth antennal segment. Concavity of propodeum well-devel-
oped over entire length, broader and more shallow. . P. rufitarsis

14. Oculo-malar space distinct, at least half the length of the tenth

antennal segment 15

Oculo-malar space very short or almost lacking. Clypeus always
much narrower than vertex. Outer orbit usually narrower than
eye in profile 17

15. Black, with partly black wings. Eyes hairy. A short but distinct

carina between occiput and middle part of outer orbit (not
behind vortex). A fine, sharp humeral collar. Clypeus about as

high as wide P. ignobilis

Russet, infuscate or blackish, with many pale (yellowish) mark-

280 bulletin: museum of comparative zoology

ings, particularly on the thorax (mesonotum striped). Wings
slightly tinged with yellow. No carina between occiput and
outer orbit or vertex. No humeral collar. Eyes bare. Clypeus
much wider than high. Outer orbit slightly wider than eye in
profile 16

16. Larger species; fore wing 11 to 11.5 mm. long. Oculo-malar space

about as long as tenth antennal segment, longer than the stretch
of the inner orbit contiguous to the clypeus. Clypeus somewhat

narrower P. singularis

Smaller species; fore wing 8.5 to 9.5 mm. long. Oculo-malar space
about half the length of the tenth antennal segment, shorter
than the stretch of the inner orbit contiguous to the clypeus.
Clypeus very wide ' P. emaciata

17. Large species (15 to 18 mm. long); black with extensive pale

markings; mesonotum always with two longitudinal pale stripes.
• Upper outer orbit separated from occiput by a strong carina,

which however does not continue behind vertex. Propodeum

with at least traces of striation. (P. striata and P. liliacea:

see couplet 3)
If pale longitudinal stripes are present on mesonotum, the species

is much smaller. Propodeum without traces of striae 18

18. Thorax shiny, with many distinct punctures. A distinct but short

oculo-malar space. Eyes with scattered hairs. Longitudinal
groove of propodeum deep. Russet, with yellowish markings
and infuscate areas. Wings nearly hyaline, very slightly

yellowish P. gorytoides

Thorax practically impunctate 19

19. Humeral margin of pronotum barely curved, raised on the sides

into a sharp, low collar which projects as fairly distinct rounded
angles. Groove of propodeum weak or obsolete ; propodeum with
erect hairs. No carina between occiput and vertex. Body 12

to 13 mm. long P. rejecta

Humeral margin of pronotum distinctly curved, either not raised
or with a low collar which never forms angles at the sides .... 20

20. Sides of pronotum with a distinct though low humeral collar.

Upper outer orbits and vertex not separated by a carina from
occiput. Propodeum distinctly grooved, with erect hairs. Body

9 to 12 mm. long 21

Humeral collar barely indicated or absent. In doubtful cases, with
a fine carina between occiput and vertex 22

21. Thorax slender, flattened dorsally, particularly on posterior por-

bequaert: social vespidae of the guianas 281

tion of mesonotum and on scutellum. Scutellum not grooved.

Eyes with fine, scattered hairs P. procellosa

Thorax more thickset, not flattened dorsally; scutellum convex,

with a longitudinal groove. Eyes bare P. catillifex

22. Upper outer orbits and vertex separated from occiput by a very

fine carina, somewhat interrupted medially. Body 8 to 11 mm.

long P. occidentalis

Upper outer orbits and vertex not separated from occiput by a

carina. Body 7 to 8.5 mm. long P. bistriata

Polybia lugubris H. de Saussure (1854) was originally described as
from Guiana, but apparently by error. The species has only been taken
definitely in southern Brazil and is unlikely to occur in the Guianas.
It is not included in the foregoing key.

Polybia ujhelyii Ducke (1909) is known from the Lower Amazon
(Brazil) and French Guiana. I have seen it from Bolivia.

Polybia signata Ducke (1910) is known from French Guiana, Brazil
and Panama.

Polybia a finis R. du Buysson (1908) was described from French
Guiana. Ducke records it also from Brazil and Ecuador. I do not
know this species.

Polybia emaciata Lucas (1879). This is in my opinion the oldest
valid, recognizable name for the well-known South American wasp
which builds a nest of clay with a circular entrance. It is generally
called Polybia fasciata H. de Saussure (1854), which was, however, a
misidentification of Polybia fasciata Lepeletier (1836). Lepeletier's
fasciata was certainly not the Polybia with the clay nest, the size given
being much too large (7 French lines = 15 mm.). I regard P. fasciata
Lepeletier, Vespa fasciata Olivier (1791) and Vespa fulvo-fasciata
Degeer (1773) as one and the same species of Stelopolybia. Polybia
caementaria Ducke (1904) is a synonym of P. emaciata. The species is
known from Dutch Guiana, French Guiana, Venezuela, Colombia,
Brazil, Peru, Bolivia, Panama, Costa Rica, Guatemala and the Repub-
lic of Honduras.

Polybia gorytoides Fox (1898)

British Guiana: west bank of the Demerara River; Bartica. Also
known from Brazil (Oyapoc; Amazon Basin; Chapada), Dutch Guiana
(Kwakoegron, Saramacca River), Bolivia (Buena vista, Dept. Sta.
Cruz) and Colombia (Villavicencio, Int. del Meta).

282 bulletin: museum of comparative zoology

Polybia liliacea (Fabricius, 1804)

P. liliacea and P. striata (Fabricius) (Syn. : P. syncophanta Gribodo)
are closely allied species, with the same type of coloration, namely
black, with extensive pale markings on thorax and abdomen and par-
ticularly with two broad longitudinal pale stripes on mesonotum.
These two species are structurally distinct and, as Dr. C. Geijskes
writes me, differ in the nesting habits.

In P. liliacea the striation and puncturation of the propodeum are
weak or obsolete, the concavity showing only faint traces of striae.
The postscutellum is rather strongly swollen. The first abdominal
tergite is short and much swollen, the swelling being rather abrupt in
profile, the basal stalk-like portion well-defined and at most one-third
of the total length of the tergite. Usually the color markings are very
pale yellow or ivory-yellow, the longitudinal stripes of the mesonotum
often coalescent before the scutellum; all or most of the abdominal
segments bear apical bands. This type of coloration was correctly
described by Fabricius as follows: "Caput cum antennis nigrum.
Thorax ater limbo lineolisque duabus crassis, antice abbreviatis,
postice coeuntibus, scutello lineolisque duabus sub scutello flavis.
Abdomen nigrum segmentorum marginibus flavis. Pedes nigri."

According to Dr. Geijskes, the nest of P. liliacea is very long (1.5
m.) and sausage-shaped, truncate at the lower end. It was partly
described by de Saussure (1858, Et. Fam. Vesp., 2, pp. cx-cxi) and
H. Lucas (1867).

British Guiana: Warina; Oronoque River; Georgetown; Berbice
Savannahs; west bank of Demerara River; Kartabo; Bartica; source
of Essequibo River. It is known also from French Guiana, Dutch
Guiana, Brazil, Venezuela, Colombia, Peru, Bolivia, and Ecuador.

Polybia striata (Fabricius, 1787)

Polybia sycophania Gribodo (1891) I regard as a synonym of
Fabricius' Vespa striata. Those who, like Ducke, refuse to recognize
this wasp in Fabricius' description, will have to use Gribodo's name.

In P. striata, the puncturation of the propodeum is coarse and the
striation is distinct though irregular and particularly well developed in
the concavity. The postscutellum is moderately swollen. The first
abdominal tergite is elongate and gradually widened, being slightly
swollen in profile, with the basal stalk at least one-third of the total
length and poorly defined. Usually the color markings are darker

bequaert: social vespidae of the gutanas 283

yellow, often with an orange tinge and the stripes of the mesonotum
may be free throughout. The apical fascial' of the abdomen are either
as in liliacea, or much reduced, sometimes almost lacking. Fabricius
evidently described one of these very dark specimens: "Caput,
abdomen, pedes nigra, immaculata. Thorax niger margine antico
tenuissime, lineis duabus dorsalibus, lateribus baseos obliquis, scutello
maculisque sub scutello flavis. Alae albae costa fusca." It should be
noted that some specimens with the structural characters of striata
are colored exactly like liliacea, so that the color is not a reliable char-
acter. On the other hand, I have seen no specimens of liliacea either
with the abdomen almost black or with orange-yellow markings.

The species is as common and as widely distributed as P. liliacea.
I have seen it from British Guiana : Kartabo ;Warina. Also from Dutch
Guiana, Trinidad, Brazil, Colombia, Bolivia and Peru. It was first
described from French Guiana.

According to Dr. Geijskes, the nest of P. striata is a white spherical
structure hung up high in a tree. It is of interest that the Indians in
the interior of Dutch Guiana distinguish P. striata and P. liliacea by
name. The nest of P. striata has not yet been described.

Ducke (1910) includes P. striata and P. liliacea in his key among the
species without striation on the propodeum. As the striation is fairly
distinct, at any rate in P. striata. I have inserted these species twice in
my key.

Polybia jurixei H. de Saussure (1854)

A widely distributed species, which I have seen from British Guiana :
Kartabo. It is also known from French Guiana, Dutch Guiana, Brazil.
Bolivia, Peru, Ecuador and Colombia (Restrepo, Int. Meta)

Polybia bifasciata H. de Saussure (1854)

This species, which extends from Mexico to southern Brazil, is
rather variable in color. In a recent paper (1943, Jl. New York Ent.
Soc, 50, for 1942, pp. 300-303), I recognize seven forms by name.
Only two of these have been taken in the Guianas thus far. Typical
P. bifasciata I have seen from British Guiana (Kartabo; Demerara
River), as well as from eastern Peru. The var. quadricincta H. de
Saussure (1854) is the most common form of the species. It occurs in
British Guiana: Bartica; Kartabo; source of Essequibo River; Warina,
X. W. District. I have also seen it from French Guiana, Trinidad,
Venezuela, Colombia, Peru and Bolivia.

284 bulletin: museum of comparative zoology

Polybia sericea (Olivier 1791)

Olivier's typical form has the head black, the thorax and first
segment of the abdomen fulvous, and the remainder of the abdomen
blackish. This is a common wasp throughout most of Central and
South America, from Guatemala to Buenos Aires. It was originally
described from French Guiana. It occurs in British Guiana (Upper
Rupununi River) and I have seen it from Dutch Guiana (Paramaribo).
Rhopalidia rufithorax Lepeletier (1836), Apoica cubitalis H. de Saussure
(1854) and Polybia melanocephala Cameron (1906) were based upon
typical P. sericea.

Most specimens I have seen from British Guiana (Demerara; Onver-
wagt ; Georgetown ; Mt. Roraima ; Bartica District) belong to the form
of the species with both first and second abdominal segments fulvous.
This is Polistes nigripennis Fabricius (1804) and, if a name is needed for
it, may be called P. sericea var. nigripennis (Fabricius). I have also
seen it from Venezuela.

Polybia chrysothorax (Weber, 1801)

Polistes aurulenta Fabricius (1804) and Polybia aurichalcea H. de
Saussure (1854) are synonyms.

Common in British Guiana : Kartabo ; Kalacoon ; Demerara ; Bartica
District; Georgetown; Forest Settlement, Mazaruni River. I have
also seen it from Dutch Guiana, Colombia, Brazil, Venezuela and
Panama.

Polybia micans Ducke (1904)

1. Typical P. micans is russet to brownish-russet with more testa-
ceous areas, particularly at the apical margins of the abdominal seg-
ments and the base of the second tergite. The tegulae and legs are
russet-testaceous, the wings strongly tinged with russet. In British
Guiana it was taken at Kartabo and Bartica. I have also seen it from
French Guiana, Dutch Guiana, Brazil, Bolivia, and Ecuador. It
appears to be somewhat nocturnal and occasionally flies to light.
Polybia sericeibalteata Cameron (1906) I regard as a synonym of
typical micans.

2. P. micans var. velutina Ducke, was described by Ducke (1907)
as a distinct species, but I am unable to separate it by structural char-
acters from P. micans. It is fairly uniformly brownish-black, the tegu-
lae and legs being also of that color. There are a few testaceous blotches
on the head. The wings are more yellowish than russet. This form is

bequaert: social vespidae of the guianas 285

rather rare in British Guiana: Arakaka. It is known also from Brazil,
Bolivia, Peru, Ecuador and Colombia.

Polybia dimidiata (Olivier, 1791)

British Guiana: West bank of Demerara River; Oronoque River,
2°42'; Penal Settlement, Bartica District. I have also seen it from
Brazil, Dutch Guiana, French Guiana, Bolivia, Peru and Colombia.

The large syrphid fly, Ceriodes braueri Williston, is perfectly homeo-
chromic with this wasp.

Polybia rejecta (Fabricius, 1798)

I recognize three color forms of this common and widely distributed
wasp, but transitional specimens are frequent.

1. Typical P. rejecta has the head, thorax, and first abdominal seg-
ment black, with very few pale yellow markings (hind margin of
postscutellum and narrow apical margin of first tergite) ; the remainder
of the abdomen dull brick-red, with or without narrow, pale margins
(sometimes ventrally only). Sometimes the red turns fuscous or black-
ish. Common in British Guiana: Mt. Everard; Arakaka; Amatuk;
source of Essequibo River; Tumatumari, Potaro River; west bank of
Demerara River. Kartabo; Bartica; Rupununi; Penal Settlement,
Bartica District; Moraballi Creek, Essequibo River. I have also seen
it from Dutch Guiana, French Guiana, Trinidad, Venezuela, Brazil
and Peru. Polybia bicolor F. Smith (1857) and Eumenes impunctus
Provancher (1888) are synonyms.

Polybia rejecta race javaryensis Cameron (1906) is a species of
Mischocyttarus.

2. Var. belizensis Cameron (1906). Entire body black, the pale
markings of the thorax as in the typical form; abdomen dorsally with
a narrow apical margin on first segment only. This is the common
form in Central America, where I have seen it from British Honduras,
Guatemala, the Republic of Honduras, Costa Rica, and Panama; but
it occurs also in Colombia, Brazil and Peru.

3. Var. litoralis Zavattari (1905). Like var. belizensis, but all or most
of the abdominal tergites with rather broad pale apical margins.
Described from Ecuador. I have seen it from Colombia, where it is
common; but Ducke's locality "Bogota" is due to an erroneous label,
as no member of the subfamily Polybiinae occurs in the vicinity of
that city.

286 bulletin: museum of comparative zoology

Polybia occidentalis (Olivier, 1791)

The most common social wasp of tropical America, from Mexico
(northward to Nuevo Leon and Tamaulipas) to northern Argentina
and Uruguay. In the Antilles proper it is known only from St. Vincent
and Grenada. It is exceedingly variable in size, color and even the
shape of the first abdominal segment, but all of the forms intergrade.
The following seem to be sufficiently distinct to deserve names.

1. Typical P. occidentalis is black, with few or many, reduced or
more extensive pale yellow or whitish markings, fairly evenly dis-
tributed over head, thorax and abdomen; most or all of the abdominal
tergites have pale apical margins; mesonotum usually unstriped.
Vespa pygmaea Fabricius (1793), Polybia pygmaea var. minor Moebius
(1856), P. pygmaea var. major Moebius (1857), P. albo-picta F. Smith
(1857), P. bohemani Holmgren (1808) and Eumenes ductus Provancher
(1888) were based on some of the variantswhich I include undertypical
occidentalis. If desired, small specimens, with more slender first tergite
and few white markings, might be called var. pygmaea (Fabricius).

This form extends over practically the entire range of the species
and is common in the Guianas. British Guiana: Kartabo; Bartica
District; Demerara; Oko River (tributary of Cuyuni River); Upper
Rupununi.

Polybia fastidiosuscula var. nigriceps Zavattari (1906) appears to be
based upon specimens of P. occidentalis much like the typical form, but
with the head entirely black and traces of pale longitudinal stripes on
the mesonotum. I have seen such specimens occasionally from colonies
of otherwise typical occidentalis and cannot regard them as more than
individual variants.

Myraptera elegans Curtis (1844) was somewhat more extensively
bright yellow, particularly on the head, and had two yellow lines on
the mesonotum. It is doubtful whether it should be given varietal
status. It is clearly transitional to var. fiavifrons.

2. P. occidentalis var. fiavifrons F. Smith (1857) has the head almost
entirely yellow (usually with an orange tinge) and is also extensively
yellow on thorax and abdomen. In particular the mesonotum bears
two broad longitudinal stripes, often fused before the scutellum.
Polybia saussurei Holmgren (1808) is not separable, according to the
types I saw in Stockholm. This form is homeochromic with the
darker specimens of P. bistriata (Fabricius) ( = occodoma de Saussure);
but the structural characters given in the key readily separate the two
species.

bequaert: social vespidae of the guianas 287

I have seen the var. flavifrons from Ecuador (Guayaquil), Peru
(Trujillo) and Brazil (Coary, Amazonas; Lassance, Minas Geraes;
Tres Lagoas, Matto Grosso; etc.).

3. Var. parvula (Fabricius, 1S()4i was originally described as almost
wholly black, with only a narrow pale apical margin on the first
abdominal segment. Many transitions connect this form with typical
ocddentalis. I am unable to separate from it P. occidentalis var.
diguetana R. du Buysson (1905).

The var. parvula often occurs in the same districts as the typical
form, but is relatively rare in British Guiana: Demerara ; Kartabo;
Kaieteur, Arakaka. I have also seen it from Mexico, Guatemala,
Costa Rica, Panama, Colombia, Venezuela, French Guiana, Dutch
Guiana, and Brazil.

4. Var. juruana R. v. Ihering (1904) is remarkable for the extension
of the yellow markings on the abdomen, the second tergite being mostly
of that color. The mesonotum, however, has no yellow stripes. I have
seen it from Brazil (Puerto America, Rio Putumayo), Bolivia, Peru,
and Venezuela; it was described from the Jurua River, Brazil.

Specimens from Aquidauana (Matto Grosso), Brazil, in which the
apical yellow margin of tergite 2 is considerably widened on the sides,
but narrow in the middle, connect var. juruana with typical occident-
alis.

5. Var. scutellaris (White, 1S41) differs from typical occidentalis only
in the yellow being restricted to scutellum and postscutellum, which
are partly or wholly of that color.

I have seen it from Brazil (Bahia; Sao Paulo), Paraguay (Villarrica),
northern Argentina (Entre Rios) and Uruguay (Montevideo).

6. Var. spilonota Cameron (1904) has not only the scutellum and
postscutellum mostly dark yellow, often with an orange tinge, but also
a large orange-yellow spot (sometimes divided) on the propodeum, as
well as other markings on head, thorax and abdomen; often, but not
always, the mesonotum bears a prescutellar spot, divided anteriorly
into two prongs; the markings of the head are rather small.

It appears to be common in Nicaragua (San Marcos, topo-
types taken with the holotype, but probably not seen by Cameron;
Managua; Chinandega; Granada). I have also seen many specimens
from Mexico, Costa Rica and Panama which provide all transitions
between typical occidentalis and var. spilonota, sometimes apparently
in the same nest.

7. Var. ruficeps Schrottky (1902) is very distinct, the head being
entirelv or mostlv brick-red to dull red, with or without vellow mark-

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ings. The remainder of the body is much marked with yellow, but
without mesonotal stripes and with the apical margins of the tergites
complete, narrowed or interrupted medially.

It is common in northern Argentina, but occurs also in southern
Brazil (Corumba, Matto Grosso), Paraguay, and Bolivia (Buena
Vista de Sta. Cruz).

Polybia procellosa Zavattari (1906)

Typical procellosa, known from Ecuador only, is mostly ferruginous-
brown, variegated with black and yellowish markings, the legs yellow-
ish-russet, the wings gray with yellowish tinge.

The var. dubitata Ducke (1910) is black with a few whitish markings,
the legs black, the wings grayish without yellowish tinge. It was
originally described from the Amazon Basin in Brazil ; but I have seen
it from British Guiana (Kurupung) and Peru.

Polybia bistriata (Fabricius, 1804)

The species generally called Polybia oecodoma H. de Saussure (1854),
is Polistes bistriata Fabricius (1804), according to Fabricius' types. I
regard it as structurally distinct, not as a color variety of Polybia
occidentalis. The characters given in my key show that it is more
closely related to P. catillifex. Cameron's Polybia brunneiceps (1912),
described from British Guiana, without more definite locality, appears
to be a synonym of P. bistriata. The ground color varies from blackish-
brown to pale russet.

P. bistriata is common in British Guiana: Kartabo; Moraballi
Creek, Essequibo River; Rockstone, Essequibo River; Demerara;
Warina, N. W. District; Kalacoon; Mt. Roraima; Penal Settlement,
Bartica District. I have also seen it from Brazil, Dutch Guiana,
Venezuela, Ecuador, Peru, and Colombia.

Polybia catillifex Moebius (1856)

British Guiana: Turesi Falls; Kamakusa; Bartica; Pakaraimo Mts.,
head of Mazaruni River. Also known from Dutch Guiana, Brazil,
Ecuador, Colombia, and Peru.

Polybia singularis Ducke (1909)

British Guiana: Bartica District; source of Essequibo River. Known
also from the Amazon Basin in Brazil, Colombia (Mapiri) and eastern
Peru.

bequaert: social vespidae of the guianas 289

As shown in the key, this is closely allied to P. cmaciata. It builds a
similar nest of clay, but with a long, narrow, slit-like (not a circular)
entrance. Such nests were described by F. Smith (1851) and H. Lukas
(1890), but without knowledge of the builder, which was first recog-
nized by Ducke (1905; erroneously referred to P. cmaciata).

POLYBIA TINCTIPENNIS Fox (1898)

The typical color form of this species is known from southern Brazil,
eastern Peru, Honduras and Panama. In British Guiana (Kamakusa;
Kartabo; source of Essequibo River) it is replaced by the var. ncbulosa
J. Bequaert (1943), with uniformly infuscated wings. This form occurs
in Trinidad and probably also in the Amazon Basin of Brazil.

Polybia rufitarsis Ducke (1904)

I have seen the typical form of this species from British Guiana:
Moraballi Creek, Essequibo River. It is also known from Brazil.

Polybia ignobilis (Haliday, 1836)

The involved synonymy of this common South American wasp will
be discussed elsewhere. Polybia atra de Saussure (1854; not Vcspa
atra Olivier, 1791) and Polybia nigra de Saussure (1858) are synonyms.

I have seen a few specimens from British Guiana: Mt. Roraima. It
is known definitely also from Panama, Colombia, Venezuela, Brazil,
Paraguay, Bolivia, Peru and Argentina.

Apoica Lepeletier (1836)

There is only one structural species of Apoica, extremely variable in
color.

Apoica pallida (Olivier, 1791)

A. pallida is a nocturnal wasp, frequently attracted by light. I have
recently examined several thousands of these wasps. The conclusions
reached, as to color variation and distribution, will be embodied in a
paper to be published elsewhere. I am able to distinguish by name five
color forms, all of which occur in the Guianas. Although each form is
somewhat variable, there are few truly transitional specimens. The
females may be separated by means of the following key.

290 bulletin: museum of comparative zoology

1. Body testaceous, pale fulvous, or darker brown, either uniformly

so or with part to most of the abdomen paler than head and
thorax. No or very few pale yellow markings (none on head and
mesonotum). Wings more or less infuscated, often blackish. . . .

var. thoracica

Body testaceous to pale fulvous, with many yellowish markings or

extensively whitish or yellow 2

2. Abdomen (except base of tergite 1) dorsally pale yellow or nearly

white, often with silvery pubescence ("frosty"); venter either
yellow or pale yellow or fulvous. Head and thorax pale fulvous
or testaceous, usually spotted with yellow. Wings subhyaline,

brownish-russet along the costa var. pallens

Dorsum of abdomen not mostly pale yellow nor silvery 3

3. All abdominal tergites with a pale yellow, narrow or wide, apical

margin; sixth tergite entirely yellow. Head and thorax spotted
with yellow (mesonotum often with two or four yellow stripes).

Wings subhyaline, slightly pale russet along the costa

var. arborea
Abdominal tergites not all margined with pale yellow 4

4. Abdomen pale fulvous (except first tergite), with apex of first tergite

and broad base of second tergite pale yellow. Head and thorax
blackish without yellow spots (not stripes on mesonotum). Wings

strongly infuscated, nearly violaceous-black var. albi macula

Pale fulvous to darker brown. Sometimes with apex of first tergite
spotted with yellow ; sixth tergite as a rule mostly yellow. Head
and thorax with many pale yellow spots. Mesonotum with longi-
tudinal stripes (at least traces). Wings slightly russet, darker
along costa typical pallida

1. Typical A. pallida. — Syn.: Vesta pallida Olivier, 1791; Apoica
lineolata Lepeletier, 1836; Polistcs translucida Spinola, 1851; Apoica
bilineolata "Lepeletier" H. de Saussure, 1854; Apoica lineata "Lepele-
tier" R. du Buysson, 1906.

A common form in British Guiana: Rockstone, Essequibo River;
Mackenzie, Demerara River; Bartica; Kartabo; Upper Rupununi
River; Demerara River; Tumatumari, Potaro River; Torani Ranch,
Berbice River; Shudihar River; Wismar; Penal Settlement, Mazaruni
River; Kamakusa. Also known from British Honduras, Trinidad,
Dutch Guiana, Brazil, Ecuador and Peru.

2. A. pallida var. thoracica R. du Buysson, 1906.

bequaert: social vespidae of the guianas 291

This form is probably as common as typical pallida. British Guiana :
Mackenzie, Demerara River; Tumatumari, Potaro River; source of
Essequibo River; Kuyuwini River; Shudihar River. It is known also
from Costa Rica, Panama, Colombia, Venezuela, French Guiana,
Dutch Guiana, Brazil, Bolivia and Peru.

3. A. pallida var. pallens (Fabricius). — Syn.: Polistes pallens
Fabricius, 1804; Apoica pallida Lepeletier, 1836.

Very common in British Guiana: Kamakusa; Shudihar River;
Oronoque River, 2° 42'; Kartabo; Tumatumari, Potaro River; Kuyu-
wini River; Georgetown ; source of Essequibo River. I have seen it also
from British Honduras, Guatemala, Nicaragua, Costa Rica, Panama,
Colombia, Venezuela, Trinidad, Dutch Guiana, French Guiana, Brazil,
Paraguay, Bolivia, Peru and Ecuador. It is reported from southern
Mexico.

4. A. pallida var. arborca H. de Saussure. — Syn. : Apoica arborea
H. de Saussure, 1854.

British Guiana: Kartabo; Torani Ranch, Berbice River; Shudihar
River; Tumatumari, Potaro River; Mazaruni River. Also known from
Dutch Guiana, French Guiana, Brazil, Bolivia, eastern Peru and
Ecuador.

5. A. pallida var. albimacula (Fabricius). — Syn.: Polistes albima-
cula Fabricius, 1804; Polistes albimaculata "Fabricius" H. de Saussure,
1854.

A rare form known only from Dutch Guiana and British Guiana:
Shudihar River; Kartabo; Pakaraimo Mis. at the headwaters of the
Mazaruni River.

Stelopolybia Ducke (1910)
(Including Gymnopolybia Ducke, 1914)

Ducke (1914, Zool. Lahrb., Abt. Syst., 36, pp. 317 and 327) divided
his genus Stelopolybia into two groups, restricting the earlier name to
the species which build nests in the open, but with the combs enclosed
in a paper envelope. The species building uncovered combs inside
some natural cavity, he separated as Gymnopolybia. Although he
mentioned some structural differences between these two biological
groups, he could find none that divided the species either consistently
or naturally.

Polistes angulata Fabricius (1804) was selected as the type of Stelo-
polybia by R. Lucas (1912, Arch. f. Naturgesch., 77, Bd. 4, Heft 1,

292 bulletin: museum of comparative zoology

p. 210).1 Unfortunately this is one of the species with uncovered
combs which Ducke later placed in Gymnopolybia. O. W. Richards
(1943, Proc. Ent. Soc. London, ser. B, 12, pts. 3-4, p. 45) has now
chosen Polybia vulgaris Ducke (1904), which I regard as a synonym
of Vespa fulvofasciata Degeer (1773), as the type of Gymnopolybia.
This is also one of the species with uncovered combs. It thus appears
that the designated genotypes of Stelopolybia and Gymnopolybia are
strictly congeneric, making the two names synonyms and leaving the
group of species building combs inside a paper envelope without a
distinct name. I feel, however, that the distinction between the two
groups is purely biological. The morphological differences assigned to
them by Ducke and Richards are either inadequate, showing transi-
tions or not applying to all the species of each group, or else of very
secondary importance in the general classification of the Polybiinae.
For this reason I do not propose a new name for the species which build
combs within an envelope.

H. de Saussure (1854) divided his subgenus Polybia, s. sir. into a
number of divisions : Alpha, Iota, Phi, Mu, Kappa and Omega. Accord-
ing to F. J. Griffin (1939, Jl. Soc. Bibl. Nat. Hist., 1, pp. 211-212),
these were all published the same year (1854) and antedate de Saus-
sure's use of some of them for groups of solitary wasps. Only Phi
and Mu contained species now placed in Stelopolybia, in addition to
others. In view of the divergence of opinion regarding the use of
Greek letters spelled out as generic or subgeneric names, I select types
which will make them synonyms of older names. Phi originally con-
tained 10 species, 8 of which are now placed in Stelopolybia and one in
Mischoeyttarus, one being unrecognized though probably also a Stelo-
polybia. I herewith designate as type Vespa phthisica Fabricius (1793),
now placed in Mischoeyttarus subgenus Kappa de Saussure (1854) (see
O. W. Richards, 1941, Proc. Ent. Soc. London, B, 10, pp. 125-126).
Mu comprised 11 species, 6 now placed in Polybia, 2 in Stelopolybia
and 3 in Mischoeyttarus. I herewith designate as type Myrapetra
scutellaris White (1841), now placed in Polybia and which is, more-
over, the monotype of Myrapetra White (1841).

iR. Lucas' earlier selection of P. angulata invalidates O. W. Richards' recent (1943) choice of
Polybia infernalis de Saussure as the type of Stelopolybia.

bequaert: social vespidae of the guianas 293

Key to Guiana Species

1. Dorsal area of pronotum rounded off, at most with a trace of

humeral collar 2

Pronotum with a raised humeral collar, sometimes projecting as
lateral angles. Oculo-malar space long 5

2. Small species, 10 to 12 mm. long. Eyes with rather distinct short

hairs 3

Large species, 15 to 20 mm. long. Eyes scarcely hairy. Oculo-
malar space short, but distinct 4

3. Oculo-malar space very long (about as long as ninth antennal seg-

ment). Upper plate of mesepisternum longer than high

S. cajennensis

Oculo-malar space very short (eye almost touching mandibular

condyle). Upper plate of mesepisternum nearly as long as high

S. pollens

4. Clypeus with distinct, fairly large punctures. Anterior margin of

pronotum moderately raised near the coxae. Lower outer orbit
nearly as wide as eye in profile. Larger, wing 15 to 16 mm. long

S. paracusis

Clypeus with fine punctures. Anterior margin of pronotum strongly

raised into a translucent lamella near the coxae. Lower outer

orbit narrower than eye in profile. Smaller, wing 13 to 14 mm.

long S. obidensis

5. Sides of humeral collar strongly projecting and angular. Eyes dis-

tinctly hairy 6

Sides of humeral collar rounded off, not or scarcely projecting. Eyes
with a few sparse hairs 8

6. First abdominal segment slender, longer than the combined pro-

podeum and postscutellum. Wing 14 to 15 mm. long, slightly

yellowish S. constructrix

First abdominal segment scarcely longer than the propodeum .... 7

7. Body orange-yellow with russet areas, the hind half of abdomen

black; second tergite with yellow base and apical margin. Wings

strongly russet-yellow, 17 to 18 mm. long <S. testacea

Body black rarely with a few yellow markings. Wings subhy aline,
with a yellowish tinge, about 16 mm. long S. angulata

8. Larger; wing about 14 mm. long. First abdominal segment rather

abruptly widened and bell-shaped, slightly angular at the sides
seen from above. Second tergite rather suddenly widened behind
the base S. fuho-fasciata

294 bulletin: museum of comparative zoology

Smaller; wing 9 to 11 mm. long. First abdominal segment very
gradually widened, not bell-shaped. Second tergite gradually
behind the base S. pallipes

Stelopolybia cajennensis (Fabricius, 1798)

Poly bia lignicola Ducke (1904) is a synonym.

A common species in British Guiana: Bartica; Kamakusa, Warina,
N. W. District; Kartabo; Forest Settlement, Mazaruni River. Also
known from Dutch Guiana (Saint Barbara Plain, Surinam River;
Paramaribo), French Guiana, Trinidad, Brazil, Colombia, Peru,
Ecuador, Panama, and the Republic of Honduras.

Stelopolybia pallens (Lepeletier, 1836)

The type of Rhopalidia pallens Lepeletier is now in Spinola's collec-
tion at the Turin Museum, where it was recognized by Ducke as identi-
cal with Polybia inf emails de Saussure (1854). Lepeletier 's name is
valid; he described his R. pallens as a new species, without any refer-
ence to Polistes pallens Fabricius (1804), which is not placed now in
Stelopolybia. Other synonyms are: Polistes rufina "Illiger" Erichson
(1848), Polybia ampullaria Moebius (1856); Eumenes fiavopectus
Provancher (1888); and Polybia internalis Dalla Torre (1904).

Common in British Guiana: Bartica; Kamakusa; Kaieteur; Turesi
Falls; Kartabo; Baracara, Mazaruni River; Demerara; Moraballi
Creek, Essequibo River; Courantyne River ; source of Essequibo River;
Forest Settlement, Mazaruni River; west bank of Demerara River.
Also known from French Guiana, Trinidad, Brazil, Bolivia, Peru, and
Ecuador.

Stelopolybia paraensis (Spinola, 1851)

This species occurs in three color forms.

1. Typical S. paraensis is extensively orange-yellow or pale ferrugi-
nous ventrally, blackish-brown or black dorsally with many pale yellow
markings, forming longitudinal stripes on the mesonotum and apical
fasciae on most of the tergites; base of first and second tergites also
yellowish; antennae and legs orange-russet; tegulae yellowish; wings
strongly tinged with yellow. Not rare in British Guiana: Mt. Roraima
Kalacoon; Pakaraimo Mts., head of Mazaruni River; Shudihar River.
Also known from Brazil, Peru, and Bolivia. Homeochromic with S.
obidensis.

bequaert: social vespidae of the guianas 295

2. S. paraensis var. ruficornis Ducke (1905). Mostly black; anten-
nae russet (scape darker); spots on sides of frons, outer orbits, hind
margin of pronotum, tegulae, longitudinal streaks on propodeum,
tibiae, tarsi, apical margins of some of the tergitcs and sternites, more
or less testaceous or whitish; no stripes on mesonotum. Wings as in
typical form. Originally described from the Upper Amazon, Brazil
(Japura River; Tabatinga). I have seen it from Peru (Iquitos; Rio
Tapiche). Ducke also reports it from eastern Ecuador.

Ducke records specimens from Chiriqui, Panama, transitional
between typical paraensis and ruficornis. I have seen such a specimen
from Pozuzo, Peru. It has the legs mostly yellowish and two narrow
yellowish lines on the mesonotum; otherwise it is like ruficornis.

3. S. paraensis var. (or subsp.) obscurior, new.

Female. — Body black; only the flagellum (particularly below) and
fore tibiae and tarsi somewhat russet; lower inner orbital margins
orange or dirty yellow, filling the ocular sinuses. Tegulae black. Wings
as in typical form.

Ecuador: Jatun Yucu, Rio Napo Watershed, 700 m., holotype and
paratypes (W. Clarke-Macintyre). — Colombia: Restrepo, Int. Meta,
500 m., paratype (J. Bequaert). — British Guiaxa: Mt. Roraima,
paratype (J. G. Myers); source of Essequibo River, paratypes (J.
Ogilvie). — Dutch Guiana: Brownsberg, paratypes (D. C. Geijskes).
— Holotype and paratypes at Mus. Comp. Zool., Cambridge, Mass.;
paratypes at U. S. Nat. Mus. and Amer. Mus. Nat. Hist.

Perfectly homeochromic with S. angulata, but without the humeral
angles of that species.

Stelopolybia obidensis (Ducke, 1904)

Polybia paraensis var. luctuosa W. A. Schulz (1905) is a synonym.

British Guiana: Tumatumari; Pakaraimo Mts., head of Mazaruni
River; source of Essequibo River; Kamakusa; Tukeit; Forest Settle-
ment, Mazaruni River; Turesi Falls; Kartabo; Bartica; Moraballi
Creek, Essequibo River; Penal Settlement, Bartica District. I have
seen it also from Dutch Guiana and it is known from Brazil.

Stelopolybia constructrix (de Saussure, 1854)

British Guiana: Pakaraimo Mts., head of Mazaruni River; Kama-
kusa ; Kartabo ; source of Essequibo River. Also in French Guiana and
Brazil.

296 bulletin: museum of comparative zoology

Stelopolybia testacea (Fabricius, 1804)

Common in British Guiana: Kartabo; west bank of Demerara
River; Mackenzie, Demerara River; Bartica; Pakaraimo Mts., head
of Mazaruni River; Kuyuwini River; Oko River, a tributary of the
Cuyuni River; Mt. Roraima. Also seen from French Guiana, Dutch
Guiana, Brazil, Venezuela, Bolivia and Peru.

Stelopolybia angulata (Fabricius, 1804)

The typical form of this species, without yellow markings and with
black tarsi, occurs in British Guiana: Kaieteur; Bartica; Kamakusa;
Arakaka; west bank of Demerara River; Warina, N. W. District;
Pakaraimo Mts., head of Mazaruni River; Mt. Roraima. I have seen
it also from Venezuela, Dutch Guiana, Brazil, Ecuador, Peru and
Bolivia. In Guatemala, Costa Rica, Panama and parts of Colombia it
is replaced by a closely allied species, S. panamensis (Cameron), in
which the humeral angles are much less prominent.

S. angulata var. angulicollis (Spinola, 1851) has the tips of the
femora and the entire tibiae and tarsi yellowish ; while the var. ornata
(Ducke, 1905) has, in addition, some pale yellow spots on the thorax.
Neither of these is known from the Guianas.

Stelopolybia fulvo-fasciata (Degeer, 1773)

I regard Polistes hectica Fabricius (1804) and Polybia vulgaris Ducke
(1904) as synonyms. Vespa ochrosticta Weber (1801) may also have
been the same. Vespa fasciata Olivier (1791) appears to be merely a
new name proposed for Vespa fulvo-fasciata Degeer. Polybia fasciata
Lepeletier (1836) was also the same wasp.

Common in British Guiana: Pakaraimo Mts., head of Mazaruni
River; Kamakusa; Warina, N. W. District; New River, a tributary of
the Courantyne; Penal Settlement, Bartica District; source of Esse-
quibo River; Rockstone, Essequibo River; Mt. Roraima; Baracara,
Mazaruni River; mouth of Meamo River. Also known from French
Guiana, Dutch Guiana, Brazil, Bolivia, Peru, Ecuador and Colombia.

Stelopolybia pallipes (Olivier, 1791)

The unwarranted emendation "pallidipes" was first used by Dalla
Torre (1894). The following names are either synonyms or based upon
color forms: Polybia anceps de Saussure (1854); Polybia lutea Ducke

bequaert: social vespidae of the guianas 297

(1904); Polybia myrmecophila Ducke (1905); Polybia festae Zavattari
(1906); Polybia pallipes var. centralis Cameron (1907); and Polybia
pallipes subsp. cuzcoensis Schrottky (1911).

1. Olivier's original description reads: "Vespa pallide testacea,
capite thoracisque dorso nigro maculatis, ahdomine fusco apice pallido.
Elle a environ cinq lignes de long [5 French lines = 12.3 mm.]. Les
antennes sont noires, avec les premiers articles d'un fauve pale en
dessous. La tete est d'un fauve pale, avec la partie superieure tachee
de noir. Le corcelet est d'un fauve pale, avec trois lignes noires sur le
dos. Le petiole est fauve pale. L'abdomen est obscur, avec la base
d'un fauve pale. Les pattes sont d'un fauve pale. Les ailes sont trans-
parentes." The French description agrees well with what Ducke
(1910) called " Stelopolybai pallidipes" and de Saussure (1854, Et.
Fam. Vesp., 2, PL XXV, fig 2) figures as "Polybia pallipes". In this
typical form the abdomen has the first tergite (or petiole) pale fulvous,
the second yellowish fulvous with infuscate or blackish apical margin,
the remaining segments blackish; there are no distinct yellow apical
fasciae. P. lirfea Ducke does not seem to differ from typical pallipes.

2. 8. pallipes var. anceps (de Saussure) has most of the abdomen
strongly infuscate or blackish, with distinct yellow apical margins at
least on tergites 2 and 3, sometimes also on 4 and 5; the yellow base of
the second tergite is, as a rule, sharply defined from the fulvous discal
area.

3. S. pallipes var. festae (Zavattari) is the extreme melanistic form
of the species, with the body blackish-brown or black and few yellow-
ish markings on head and sides of thorax (none on mesonotum or
abdomen). It is known only from Eastern Ecuador (San Jose, 1800
m.). I have not seen it.

4. S. pallipes var. cuzcoensis (Schrottky) has the«black abdomen of
var. festae; head and thorax are profusely marked with yellow, as in
the typical form. It appears to be characteristic of parts of eastern
Peru: Achinamiza; La Chorrera, Putumayo; Chanchamayo. I have
also seen it from Bolivia : Huachi, Rio Beni. It is evidently the form
with black abdomen mentioned by Ducke from the Upper Amazon:
Tabatinga; Iquitos.

5. S. pallipes var. (or subsp.) fulvanceps, new.

Female. Ground color orange to fuscous-yellow, darker on the
dorsum of the abdomen, more yellowish ventrally; head and meso-
notum with the same blackish markings as in the other forms of the
species; narrow base of second tergite and broad and well-marked apical
margins of tergites 2 to 5 bright yellow; tergite 6 mostly yellow.

298 bulletin: museum of comparative zoology

Wings and legs as usual. This form combines the fulvous-orange
abdomen of var. myrmecophila with the abdominal bands of var.
anccps.

Colombia: Rio Frio, Dept. Magdalena, holotype and paratypes
(G. Salt). — Peru: Colonia Perene, paratype (J. C. Bradley); Puerto
Bermudez, Rio Pichis, paratypes (J. C. Bradley); Miriantiriani,
Camino del Pichis, paratypes (J. C. Bradley). — Holot3rpe and para-
types at Mus. Comp. Zool., Cambridge, Mass.; paratypes also at
Cornell University (Dept. of Entomology).

6. S. pallipes var. myrmecophila (Ducke) has the head and thorax
of the typical form, but the abdomen is almost unicolorous pale
yellowish-russet or orange, usually with more yellow bases of second
and third tergites. Sometimes the apical margins of some of the ter-
gites are very narrowly or faintly yellowish. P. pallipes var. centralis
Cameron is clearly the same form, as was recognized by Ducke.

Typical S. pallipes was described originally from French Guiana
(Cayenne) and I have seen it from British Guiana (Mt. Roraima),
where it appears to be uncommon. I also know it from Brazil, Para-
guay, and Peru (mouth of Rio Cotuhe)

The var. anceps is the usual form in British Guiana: Kamakusa;
Kartabo; west bank of Demerara River. I have also seen it from
Dutch Guiana, Trinidad, Brazil, Venezuela, Colombia, Panama and
Bolivia.

The var. myrmecophila also occurs in British Guiana: Source of
Essequibo River; Demerara River; Rupununi River. It is more widely
distributed than any of the other forms, as I have seen it from Guate-
mala, Honduras, Costa Rica, Panama, Colombia, Ecuador, Peru and
Brazil.

Pseudopolybia H. de Saussure (1863)

The genus was revised in 1938 (Rev. de Entomologia, 9, pp. 112-
113), but a new species is described below. Of the five species known
at present, four occur in the Guianas.

1. Mesepisternum not divided by an oblique suture. Humeral margin
of pronotum straight, bluntly ridged. First abdominal segment
longer than wide, forming a broad stalk. Small; fore wing 5.5
to 6 mm. long P. pusilla

Mesepisternum completely divided by a well-marked oblique suture
into an upper and a lower plate 2

bequaert: social vespidae of the guianas 299

2. Small; fore wing 5.5 to G.5 mm. long. First abdominal segment

longer than wide, forming a distinct stalk, gradually widened
from base to apex (seen from above). Clypeus slightly produced

medially P. langi

Larger; fore wing 11 to 12 mm. long. First abdominal segment
either cap-shaped, or, if stalk-like, not gradually widened from
base to apex. Clypeus strongly produced medially 3

3. First abdominal segment slightly longer than wide, forming a broad,

bell-shaped stalk, set off from the remainder of the abdomen

P. difficilis

First abdominal segment wider than long, cap^shaped and not set

off as a stalk from the remainder of the abdomen ... P. vespiceps

Pseudopolybia vespiceps (de Saussure, 1863) has been taken in
French Guiana and Dutch Guiana, as well as in Brazil.

Pseudopolybia pusilla (Ducke, 1904)

British Guiana: Monkey Jump, Essequibo River (Oxford Uni-
versity Exped.).

The species is also known from Brazil (Para and Oyapoc). It is
readily mistaken for a Leipomeles. It differs from L. dorsata in the
much wider clypeus and stronger humeral transverse ridge. The nest
is as yet unknown.

Pseudopolybia difficilis (Ducke, 1905)

I have seen this species from British Guiana: source of Essequibo
River. It is also known from Brazil, Peru, and Bolivia.

PsEUDOPOLYBtA langi, new species. Fig. 2

Female. Head moderately flattened, slightly wider than thorax;
seen in front, slightly wider than high; from above, rectangular with
receding hind corners, about twice as wide as long; occipital margin
nearly straight. Vertex and genae not margined by a carina behind;
gena scarcely narrower than the eye in profile. Oculo-malar space
about half the length of the fourth antennal segment. Inner orbits
about one and one-third times as far apart on vertex as at clypeus.
Ocelli rather large, in an equilateral triangle; posterior ocelli about
twice as far from eyes as from each other. Interantennal shield broad,
not set off, strongly but bluntly and evenly swollen. Antennae nearly

300

bulletin: museum of comparative zoology

twice as far apart as from eyes. Frons slightly convex. Clypeus at its
narrowest about one and two-thirds times as wide as high, irregularly
pentagonal with the upper side much the longest, contiguous to the
eyes over nearly one-half of sides; apical margins slightly produced,
ending in a very broad, obtusely rounded point. Mandible with an
oblique cutting edge of four sharp teeth, the lower three subequal, the
upper one much smaller; most of outer surface flattened or slightly
depressed. Antenna: scape short and rather thick, scarcely curved;

Fig. 2. Pseudopolybia langi J. Bequaert. Female. A, wings; B, head in
front view; C, maxillary palpus; D, labial palpus; E, body in profile.

second segment short, moderately swollen ; third less than three times
the length of second; fourth and fifth about as long as wide; sixth to
eleventh slightly wider than long; twelfth slightly longer than wide
at base, with bluntly rounded tip; flagellum about equally thick
throughout. Maxillary palpi of 6 segments. Labial palpi of 4 seg-
ments, the tip of the third with a short, heavy, erect, curved seta.
Thorax moderately elongate; in profile, about one and one-third times
as long as high ; from above, nearly twice as long as wide before tegu-
lae. Pronotum evenly rounded into a semi-circle anteriorly, the
humeral margin barely indicated by a very weak transverse, blunt

bequaert: social vespidae of the guianas 301

ridge on the sides, where it forms no angles ; dorso-lateral areas sloping;
lower anterior margin of ventrolateral areas with a vertical swelling
indented by a deep pronotal fovea. Mesopleuron moderately swollen ;
mesepisternum completely divided by an oblique suture into an upper
and a lower plate; deep mesepimeral suture ending abruptly and far
from the metapleuron. Upper sclerite of metapleuro'n ending obliquely
below, with a short extension along mesometapleural suture. Scutel-
lum slightly and evenly convex, anteriorly with a fine, raised, longi-
tudinal line which continues posteriorly as an impressed line. Post-
scutellum transversely elliptical, moderately swollen. Propodeum not
swollen, moderately slanting in profile; median concavity a broad,
shallow, longitudinal groove. First abdominal tergite narrowed,
moderately elongate, very gradually widened and thickened from base
to apex, where it is less than half as wide as second tergite, triangular
in outline from above. Second tergite from above wider than long,
gradually widened at base ; in profile, slightly and evenly convex. Mid
tibiae with two spurs. Claws symmetrical, unarmed. Venation:
second cubital cell much higher than wide; third cubital longer than
wide, rectangular, about as long on cubitus as on radius; radial cell
very long, acute; basal vein ending in subcosta close to the large
stigma.

Smooth and moderately shiny, without appreciable punctures or
other sculpture. Pubescence sparse and short. Eyes with many dis-
tinct erect hairs.

Head, thorax and legs pale yellowish, with a few fuscous areas on
frons, vertex, occiput, pronotum, mesepimeral suture, propodeum
(concavity and extreme sides), hind portion of postscutellum, and
tibiae. Antennae fuscous above, russet below. Teeth of mandibles
fuscous. Mesonotum fuscous, with a pair of broad median yellow
stripes and a shorter yellow streak on each side near the tegula.
Abdomen pale fuscous to light ferruginous, more or less yellowish at
the base of the second tergite. Wings hyaline throughout; veins pale
fuscous; stigma translucent medially.

Length (h.+th.+t.l+2): 5.5 mm.; of fore wing, 5.6 mm.

British Guiana: Kamakusa, female holotype and paratypes from
one nest (Herbert Lang). Holotype and paratypes at Mus. Comp.
Zool., Cambridge, Mass.; paratypes also at Am. Mus. Nat. Hist, and
U. S. Nat. Mus.

The characteristic thick seta of the labial palpus is shorter than in
some other species of the genus and, being of a pale color, is more
difficult to see.

302 bulletin: museum of comparative zoology

Parachartergus R. v. Ihering (1904)

This genus was revised by me in 1938 (Rev. de Entomologia, 9, pp.
105-112). Unfortunately I called it there Chartergus, in the belief that
Ashmead (1902) was the first to select a genotype for that name. The
late Miss Sandhouse informed me that Emile Blanchard in 1840
(Hist. Nat. Ins., 3, Orth. Nevr. Hem. Hym. Lep. Dipt., p. 395) had
designated Vespa nidulans Fabricius (1793), a synonym of Vespa
chartaria Olivier (1791), as the type of Chartergus Lepeletier. The
name Parachartergus must therefore be used for the genus which I had
called Chartergus.

Key to Guiana Species

1. Vertex and cheeks not separated from the occiput by a carina.

Humeral margin of pronotum erect, not overlapping nortouching

the vertex P. frontalis

Vertex and cheeks separated from the occiput by a sharp carina.

Humeral margin of pronotum slanting, touching or overlapping

. the vertex 2

2. Vertex on each side, near the upper inner orbit, with a slightly

raised, impunctate area. Smaller; fore wing 8 mm. long

P. smithii
Vertex without impunctate lateral areas 3

3. Body thickset, black. Humeral margin of pronotum low, ridge-

like, opaque. Wings black, with the apical portion either some-
what paler or decidedly whitish. Fore wing 9 to 12 mm. long. .4
Humeral margin of pronotum raised into a high, sometimes trans-
lucent lamella. Wings not black with paler or whitish tips.
Fore wing 6.5 to 8 mm. long 5

4. Head and thorax with distinct, long erect hairs P. apicalis

Head and thorax covered with a grayish bloom, the erect hairs

restricted to clypeus, postscutellum and propodeum

P. fratemus

5. Body lengthened, black or ferruginous. Thorax over one and a half

times as long as wide seen from above P. fulgidipennis

Body short, thickset, mostly testaceous or russet. Thorax about

one and one-third times as long as wide seen from above

P. eolobopterus

Parachartergus eolobopterus (Weber, 1801) was taken in French
Guiana and is known also from Colombia, Venezuela, and Trinidad.

bequaert: social vespidae of the guianas 303

Parachartergus frontalis (Fabricius, 1S04), not yet reported from the
Guianas, is known from the adjoining Oyapoc district of Brazil and
Venezuela.

Parachartergus apicalis (Fabricius, 1804), frequently recorded from
the Guianas, possibly does not occur there, as it is often confused with
P.fraternus, which I regard as a distinct species.

Paraciiartergus fraterxus (Gribodo, 1S91)

1 . The typical form of this species has the tips of the wings whitish
with pale veins, being colored exactly like P. apicalis. I have seen it
from Dutch Guiana (Paramaribo).

2. In the var. concolor Gribodo (1891), the tips of the wings are
somewhat paler but not whitish and have fuscous veins. I have seen
it from British Guiana (Hepseba, Courantyne River; Arakaka),
French Guiana, Brazil, Trinidad, Colombia, and Panama. It was
first described from Venezuela.

Parachartergus fulgidipennis (H. de Saussure, 1854)

I have recognized by name five color forms of this species, two of
which occur in the Guianas.

1. The var. griseus Fox (1898) has the thorax and abdomen black,
the abdomen rather dull with many long hairs, and the head partly
pale yellow. The wings are subhyaline, broadly black along the costal
margin, without distinct cream-colored transverse patch. Charter gus
trichiosomus Cameron (1912), described from British Guiana, is a
synonym. I have seen this form from British Guiana (Moraballi
Creek, Essequibo River), Brazil, and Peru.

2. In the var. fasdipennis Ducke (1905) the color of the body is as
in var. griseus, but the abdomen is rather shiny, with sparse and short
hairs, and the wings have a distinct cream-colored transverse patch,
which interrupts the black streak along the costal margin. It was
originally described from Brazil; but I have seen it from British
Guiana: Ite Cattle Trail.

Parachartergus smithii (H. de Saussure, 1854)

The typical form of this species was taken in British Guiana:
Kartabo. It also occurs in British Honduras, Costa Rica, Colombia,
Ecuador, Peru, and Brazil.

304 bulletin: museum of comparative zoology

Leipomeles Moebius (1856)
This genus is monotypic.

Leipomeles dorsata (Fabricius, 1804)

W. A. Schulz (1912, Berlin. Ent. Zeitschr., LVII, p. 87) examined
the type of Polistes dorsata Fabricius and recognized that it was the
wasp described by Moebius (1856) as Leipomeles lamellaria. Polybia
nana H. de Saussure (1863) and Polybia spilogastra Cameron (1912)
are other synonyms.

British Guiana: Source of Essequibo River; Monkey Jump, Esse-
quibo River. Fabricius' type came from the Essequibo River. Also
known from Dutch Guiana, Brazil, Bolivia, Peru, Ecuador, and
Panama (Barro Colorado).

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCIV, No. 8

THE BIRD FAUNA OF THE WEST SUMATRA ISLANDS

By S. Dillon Ripley

Division of Birds
United States National Museum

With Two Plates

CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
October,; 1944 j

PUBLICATIONS
OF THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
investigations by the Staff of the Museum or of reports by spec-
ialists upon the Museum collections or explorations.

Of the Bulletin, Vols. I to XCIII, and Vol. CXIV, No. 1, 2,
3, 4, 5, 6 and 7 have appeared and of the Memoirs, Vol. I to LVI.

After 1941 no more Memoirs are to be published.

Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE

Vol. XCIV, No. 8

THE BIRD FAUNA OF THE WEST SUMATRA ISLANDS

By S. Dillon Ripley

Division of Birds
United States National Museum

With Two Plates

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

October, 1944

Xo. 8— The Bird Fauna of the West Sumatra Islands1
By S. Dillox Ripley

CONTENTS

Page

Introduction 307

Acknowledgements 308

Ornithological History 309

Geography of the islands 311

A Faunal List of the birds of the islands 317

Summary and Conclusions 415

Bibliography 427

INTRODUCTION

The islands of the East Indies stretching from the Asiatic mainland
down to the southeast for more than two thousand m'les 'nto the
Papuan and Australian regions, form a true paradise for the student
of faunal distribution and speciation. The position of this myriad of
islands large and small, each with its varying type and degree of geo-
graphic and geologic isolating mechanisms is highly interesting. The
equable climate, with a minimum of seasonal or spasmodic changes,
adds to the effectiveness of even the most limited geographic barriers.
Ten or fifteen miles of sea is enough to preclude interchange of genes
between two sedentary populations. As an example of this kind of
speciation the bird fauna of the west Sumatra islands offers a superla-
tive opportunity for study.

Several factors have influenced my decision to study the birds of the
west Sumatra islands. During 1939 I was able to visit Nias, the largest
island of the group, where I made a small collection for the Academy
of Natural Sciences of Philadelphia. When, early in 1942 I came to the
United States National Museum from Harvard, I found that I would
have an opportunity in Washington to make an exhaustive study of
what is the largest and finest collection of birds from these islands, that
of the late Dr. W. L. Abbott. Dr. Abbott's collection has never been
reported on as a whole, but various papers by the late Dr. Charles
Richmond, Dr. H. C. Oberholser, and the late Mr. J. H. Riley have

1 In the absence of the author in war service, the proof was kindly read by Dr. Herbert
Friedmann, Curator of Birds, U. S. National Museum. Editor.

308 bulletin: museum of comparative zoology

appeared from time to time describing more than one hundred and thirty
new species and subspecies, the types of which are now in the United
States National Museum. Thus without actually returning to the
islands, now out of the question, I have been able to accumulate re-
markably complete data on the fauna of this area.

ACKNOWLEDGEMENTS x

To the late Dr. Glover M. Allen of Harvard, my councillor and
friend, I owe a great deal for what has gone into this study. Professor
A. C. Romer has been most helpful with aid and advice. Mr. James
L. Peters has been kindness itself in rendering me every encouragement
and assistance. Dr. Ernst Mayr of the American Museum of Natural
History and Mr. Rodolphe de Schauensee of the Academy of Natural
Sciences have freely loaned specimens from their collections and have
given much valuable advice. To the authorities of the United States
National Museum, Dr. Alexander Wetmore, Dr. Herbert Friedmann,
and Mr. H. G. Deignan, I owe a tremendous debt of gratitude for their
generous help and wise counsel. Certain final questions of identifica-
tion remain unsettled. For this unfortunately there is no solution
until such time as free communication, field study, and research in
general return to their proper places in this now sadly maladjusted
world. That this will happen in good time is the earnest hope and
calm conviction of all those of us who now labor in alien and exciting
fields.

S. Dillon Ripley
Washington, D. C.
November, 1942.

> Originally presented as a thesis in partial fulfillment of the requirements for the degree of
Doctor of Philosophy at Harvard University.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 309

ORNITHOLOGICAL HISTORY

The first serious scientific observing and collecting on the west
Sumatra islands was done by Baron H. C. B. von Rosenberg who
visited Nias in 1854. So far as is known the list of 56 species of birds
published by Xieuwenhuisen and himself in 1SG3 as being found on
Nias, and again the list published by him in 1878 of 60 species of
birds noted on Nias and nearby islands are not based on actual collected
specimens but only on observations. The first actual collection of
birds was made by Signor Elio Modigliani during his exploration of
Nias from April through August, 1886. This collection of 62 species,
eight of which were new, was published on by Count Salvadori in 1886
and also by Modigliani himself in 1890. By this collection the known
avifauna of Nias was raised to 102 species.

Two collections were made among these islands in 1S91. One on
Nias was prepared by M. J. Claine and reported on by Dr. Oustalet
in 1892. Signor Modigliani meanwhile had gone to Enggano and his
very interesting material consisting of 23 species, seven of which were
new, was described by Salvadori in 1892. Another collection made on
Nias was that of Mr. W. Thomas, a missionary on the island, who
sent birds to Graf von Berlepsch. Some of these birds were described
by Salvadori and some by Berlepsch in 1895. Signor Modigliani, who
was still in the islands, now visited Sipora, and Salvadori listed his
collection from there of 34 species, 3 of which were new, in 1894.

A hard-working and able young entomologist, Mr. J.Z.Kannegieter,
visited Nias and the Batu Islands during the winter of 1895-96. Mr.
J. R. H. Neervort van der Poll, for whom Kannegieter was working,
presented his bird collection to the Leyden Museum, where the Nias
collection was reported on by Dr. J. Biittikofer. Dr. Biittikofer's list
raised the known avifauna of Nias to 128 species, of which 4 were
described as new in his paper (1896). Mr. van der Poll included in-
formation on the native names of these birds as well as notes on the
color of the soft parts of the specimens. The 355 Batu specimens,
except for two species described by Finsch (1899), remained unworked
until 1940 when the Academy of Natural Sciences of Philadelphia pub-
lished two papers by de Schauensee and Ripley, and de Schauensee.

The next collections made on the islands were those of the tireless .
Dr. W. L. Abbott who voyaged about the East Indian islands for
several years on a small schooner manned by an adventuresome and
daring Malay crew. Dr. Abbott started in the autumn of 1901 at the
northern end of the chain, visiting Simalur, Babi, Lasia, and the islands

310 bulletin: museum of comparative zoology

of the Banyak group, Bangkaru and Tuangku. A year later in the
autumn and winter of 1902, Dr. Abbott along with Mr. Charles Boden
Kloss of the Federated Malay States Museum went again to Simalur.
From there they sailed south to the Pagi Islands, the Batu Islands,
Tello, Tana Massa, and Tana Bala, and lastly Nias. Two years later
in the fall of 1904, Dr. Abbott visited Enggano. His last trip to the
group was to Nias in the spring of 1905. As a result of these compre-
hensive trips more than 1300 bird skins were presented to the United
States National Museum.

Several papers have been written on these collections. Dr. Charles
W. Richmond published the first on the Simalur trip in 1903 describ-
ing 19 new forms. Much later, Dr. Harry C. Oberholser (1919) wrote
up Dr. Abbott's second collection on Simalur describing one new form.
In these two papers there are color notes on the species, and in Dr.
Richmond's paper there are some short field notes made by Dr. Abbott.
On various occasions Dr. Oberholser, notably in his well known paper,
"Descriptions of One Hundred and Four New Species and Subspecies
of Birds from the Barussan Islands and Sumatra" (1912), and Mr.
Riley have published descriptions of new forms from the Abbott
collection, but no comprehensive paper on the collection as a whole
has ever appeared. Mr. Riley at the time of his death was working
on a faunal list of the birds of these islands. It is unfortunate that he
did not live to complete it. Needless to say it has been an inestimable
source of valuable information in the preparation of this manuscript.

The next collection of birds from these islands was that of Dr. E.
Jacobson and Jonkheer W. C. van Heurn made in 1913 on Simalur and
its adjacent islands. Dr. G. C. A, Junge published on this collection
in 1936, describing 4 new races and including many extremely valuable
field notes made by the collectors. There is also a good deal of material
about the eggs of some of these forms, which were collected and
measured.

Due to difficulties with the local population, Dr. Abbott was not
given permission by the Netherlands authorities to visit Siberut and
Sipora. Later in 1924 an expedition from the Raffles Museum, partly
financed by funds given by Dr. Abbott, visited these islands and a
duplicate set of these birds was given to the United States National
Museum. This collection was reported on by Messrs. F. N. Chasen
and C. Boden Kloss in 1926. They described 11 new forms. In 1927
Mr. Riley described 3 more forms from the same collection.

So far as can be discovered the next ornithological collecting on these
islands was not until 1937 when three collectors were in the field. Miss

Fig 1 Map "i Wat Suraatni l-lnrnf-, -.|mumj» depth contours,

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 311

Barbara Lawrence made a collection of 31 species and subspecies from
Nias for the Museum of Comparative Zoology at Cambridge, Mass.
Mr. J. J. Menden made a collection on North and South Pagi, part of
which is now at the same Museum. The third collection was that of
Dr. J. K. de Jong and a worker from Buitenzorg, Saiin, who spent two
months on Enggano. The first two collections have not been worked
up, but the Enggano birds were reported on by Dr. Junge (1938), add-
ing 5 species to the list of birds found on that island. In June 1939 I
spent a few days on Nias and secured specimens representing 45 species
and subspecies. This material has been written up by de Schauensee
and myself (1940), and four new forms were described.

Altogether some two hundred twenty species and subspecies have
been described from the islands of which the present paper recognizes
one hundred fourteen forms. The total list of the bird fauna of this
small archipelago runs to two hundred eighty species and subspecies
including migrants. With the exception of three races I have been
able to see specimens of all the endemic forms occurring in this group.
These three races are: Spizaetus cirrhotics vanheurni and Eurostopodus
macrotis jacobsoni from Simalur, and Coraeina striata kannegieteri
from Nias. Present conditions indicate that collecting will be impos-
sible on these islands for some time. However, I think that except for
the recording of more migrants and the extension of range of some
known forms the present list should stand for some time to come. I
do hope that in the future there will be an opportunity for more
biological field work to be done on the birds of these islands. Un-
doubtedly they serve as a splendid natural speciation laboratory.

GEOGRAPHY OF THE ISLANDS

The west Sumatra Islands lie in a chain for a distance of approxi-
mately six hundred miles paralleling the western coast of the island of
Sumatra. This chain runs roughly from northwest to southeast ex-
tending from Lat. 3°N„ Long. 95°35'E., to Lat. 5°30'S., Long.
102°24'E. The islands have never been carefully explored by a
geologist. (See map, fig. 1).

The original appearance of these islands is presumably due to the
same phenomena which caused the rise of the Barussan Mountains in
western Sumatra. Brouwer (The Geology of the Netherlands East
Indies, 1925, p. 2) has this to remark about these mountains:

"Sumatra lies along the axis of a great geanticline. Stretching along
the western part of the island, near to the coast are the Barissan [sic]

312 bulletin: museum of comparative zoology

Mountains which give to this side of the island a rough and rugged
character. . . . The present Barissan Mountains are associated with
late Tertiary and post-Tertiary mountain-building. Considerable vul-
canism occurred during a great part of the Tertiary period and even
yet there are many active volcanoes." Later (p. 84), he says: "The row
of islands to the west of Sumatra is known for its frequent and intense
earthquakes, but volcanoes are entirely lacking."

Professor G. A. F. Molengraaff in an interesting article ("Modern
Deep-Sea Research in the East Indian Archipelago", Geog. Journ.
57, No. 2, Feb. 1921, p. 95.) has a good deal to say about the forma-
tion of the west Sumatra Islands. His discussion of folding (p. 108)
follows :

"As soon as the upper portions of the folds which develop at a cer-
tain depth approach the Earth's surface and the majority of the rocks
under diminished pressure can no more be folded without being frac-
tured (van Hise's zone of fracture), the continuity of the strata will be
broken and the culminating portions of the elevating anticlinal axes
will be fractured and show at the surface as isolated portions or blocks,
their extent and shape being greatly dependent on the geological
structure and the differences in rigidity of the composing rocks. This
may suffice to explain why an elevating submarine ridge formed by
an anticlinal axis will appear at the surface as a row of blocks, i.e.
islands separated by deep channels.

"In the great geosynclinal area between the continents of Asia and
Australia one arc of folding belonging to the Alpine system, and known
as the Malay arc, appears to originate from the Burma arc, . . . and
can be followed . . . frcm the extreme north-western end of Sumatra,
through this island and the island of Java. . . as far as the Banda sea.

"In its western section where it borders on the Indian Ocean, as in
the central portion of the island of Sumatra, this arc now consists of
two major folds, as is illustrated by an ideal cross section (Fig. 2)
from the Indian Ocean towards the stable portion of the Sunda land.
It would show the following sequence:

1. Indian Ocean.

2. Sunda trough, first geosyncline.

3. Range of coastal islands girdling the west coast of Sumatra in-
cluding the Mentawei islands, first geoanticline.

- 4. Mentawei trough and corresponding trough-shaped depths,
second geosyncline.

5. Non-volcanic and volcanic mountain ranges of Sumatra and
Java, second geoanticline.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 313

6. Tertiary terrain, folded in late Tertiary and early Pleistocene
time, now practically stable, third geosyncline.

7. Stable Sunda land including the Sunda shelf."

wsw. />/ ^^ ^ *£A LCVEL ENE

1. 2. 3.4. 5. <o. 7.

Fig. 2. Schematic representation of late Tertiary, Pleistocene, and post-
Pleistocene crustal movements in the western section of the Malay geosynclinal
area. (After Molengraaff) The numbers refer to those listed above.

The northern group of islands consists of Simalur and its outliers.
Simalur is about fifty-five miles long, varying in width from five to
fourteen miles. It lies about sixty-five miles from the coast of Sumatra.
It is a rocky well-wooded island, the main mountain of which, Sibalur,
rises to 625 metres in height. Simalur is thinly populated and there are
few gardens or clearings. Besides raising sago, coconuts and vegetables
for their own subsistance, the main industry of the people is raising
water buffalo. The principal village is Sinabang in the southeastern
part of the island.

Simalur is outside the two hundred metre line which curves out from
the Sumatran coastline and envelops the Banyak islands to the south-
east. Simalur lies on its own two hundred meter bank which encloses
several nearby islands. Northwest lie the Kokos or Sa Laut Islands,
two small flat islets which are the most northerly extensions of the
entire archipelago. These two islets are two miles by one and a half
miles, and one thousand yards in diameter respectively. They are
connected with Simalur twenty-five miles to the south east by a sub-
terranean bank less than thirty-six metres in depth. They are mainly
covered with coconut trees and the only collectors to visit them were
Jacobson and van Heurn who spent a day there, August 21, 1913.
They found two forms on Kokos, Caloenas nicobarica nicobarica and
Eudynamis scolopacea simalurensis. Presumably other birds, particu-
larly migrant shore birds, occur there from time to time.

The only other islets which have been collected on in the immediate
vicinity of Simalur are Djawi Djawi, Lugu, and Pulu Pandjang in
Sinabang harbor, and Siumat, six miles off shore in an easterly di-
rection. Siumat is about two and a half miles long with some original
forest as well as coconut palms. Both Dr. Abbott and Jacobson and
van Heurn collected there getting three common species of small island

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birds: Sterna dougalli bangsi, Chalcophaps indica indica, and Halixetus
leucogaster.

South of Simalur by about fourteen miles are two small islands
separated from each other by a strait one and one quarter miles wide.
They are sometimes called the Tapah islands. Both are small low coral
islands covered with original forest and uninhabited. Lasia, the north-
ernmost is smaller, less than two miles in length, and surrounded by
fringing reefs. Babi, the larger, is about seven miles in diameter and
nearly round. Both islands lie on a shelf separated from the neighbor-
ing islands to the north, east, and south by depths greater than two
hundred meters. The only collector to visit Babi and Lasia has been
Dr. Abbott. Together these two small islands have three endemic
races not found on any of the other islands in the group. These are :
Psittacula alexandri major, Coracina striata babiensis, and Hypothymis
azurea abbotti.

The Banyak Islands lie about thirty-eight miles off the coast of
Sumatra, and forty-five miles south-east of Simalur. Bangkaru, the
most westerly, is thirty miles east of Babi. The principal islands of
the more than fifty islets and reefs in the group are : Tuangku, seven-
teen miles long by five miles wide, Bangkaru with an area of twenty
square miles, and Ujung Batu, a small narrow island. These islands
are decidedly hilly and well forested. A mountain on Bangkaru,
Amintolan, reaches a height of three hundred three meters. There is
a hill at the south end of the island. In between the land is very low
so that at any distance the appearance presented is that of two islands.
Tuangku has a mountain of three hundred thirteen meters on the north
coast. The eastern coast is low and covered with mangroves. There is
a small population on these islands of which more than half (536 in
1911) live in Tuangku.

The only collections from the Banyak islands have been those of
Dr. Abbott who visited Tuangku and Bangkaru. Richmond (1903,
p. 485) notes that more species were seen on Tuangku than Dr. Abbott
had found on Simalur. However, Abbott noted that "no large parrots,
hornbills, or barbets were seen or heard, and no drongos or orioles
were noticed."

The largest island of the group is Nias which is seventy miles long
and from twelve to twenty-two broad. It lies twenty-eight miles south
of the Banyak islands and about fifty-six miles south west of Ujung
Singkel, a projecting point of the Sumatra coastline. Nias lies on its
own two hundred meter shelf. The nearest point of contact is with
that part of the Sumatran shelf which extends south of the Banyak

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 315

islands. The elevation of Nias reaches a height of eight, hundred eighty-
six meters. In general the central part of the island is rather high
forming an irregular plateau which extends down on all sides gradually
to the coast. The general impression is of a drowned island with an
irregular sloping outline surrounded by outlying submerged rocks and
coral reefs in all directions. Further out from the shore are many
small islands such as the Hinako group which lie on a ten fathom bank
extending out from the west coast of Nias. Presumably most of these
islands have been connected with Nias at one time or another.

Nias has the largest population of any of the islands numbering
about one hundred thousand at the last census. The main centers
are in the southern part of the island where there are many large
villages. This southern part of the island has been rather heavily
deforested with the result that original forest is now found only in a few
places on the hill tops. The northern part of the island on the other
hand is populated by a somewhat less aggressive people whose culture
apparently does not demand the construction of the giant houses found
to the south.

In this part of the island much original forest remains and there are
presumably several new records to be added to the bird list of the island
as many fewer collectors have worked here. Nias has been by far the
best collected of all the islands due to its size and accessibility. Until
1942 it was the only one of the islands which was visited regularly by
an inter-island steamer sailing from Sibolga to Goenong Sitoli every
two weeks. The main trade of the island was in pigs which were much
in demand in the Chinese-populated centers on Sumatra.

Southeast of Nias by about forty-five miles lie the Batu Islands.
These consist of three large islands, Pini, Tana Massa and Tana Bala
surrounded by more than twenty smaller ones. Besides these three
large islands, Tello and Lago are the only islands that have been visited
by collectors. Tana Massa is twenty -seven miles long by five miles
broad. Tana Bala is twenty-two miles long by seven broad. Pini is
twenty miles long and six broad. All these islands are rather low and
without distinguishing features. The population is small and the origi-
nal forest largely remains. Tello, less than two miles long, is the seat
of administration and the most populated island in the group. Lago is
a small coconut covered islet three miles northeast of Tana Massa.
Most of these islands are surrounded by reefs sometimes five or six
miles out in all directions. The whole group lies within the two hundred
meter line which curves out from the Sumatra mainland and continues
southeast in a narrow belt inclosing the islands below the Batu group.

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The name for those islands to the south of Tana Bala is Mentawi.
This includes Siberut, Sipora, North and South Pagi, and the adjacent
small islands. North and South Pagi by themselves are called the
Pagi or Pagai islands. All four of these islands are inhabited, hilly,
and evidently of volcanic formation. Although these islands lie on a
relatively shallow bank connected with Sumatra to the north, there is
a definite gap directly to the east towards the nearest part of the Su-
matran mainland. Here there is the deep Mentawi basin reaching a
depth of over sixteen hundred meters. Thus except for the one link to
the north there is no evidence to indicate that the Mentawi group has
ever been connected with Sumatra.

Siberut, thirty miles to the south of Tana Bala, is sixty miles long by
fifteen to twenty-four miles in breadth. It is heavily wooded with ex-
tensive marshes along the coast. There are about eight thousand in-
habitants at an extremely primitive cultural level. The island is of all
the group perhaps the least known and least explored. The only
collecting of birds and mammals has been that of Messrs. Kloss and
Smedley in 1924.

Sipora is a small densely overgrown island about thirty-three miles
long. The low-lying land is extremely marshy, so much so that in
former years the best communication into the interior was by canoe.
The highest point on the island is three hundred thirteen meters.
Sioban, the main village is on the eastern shore and is a small copra
shipping port.

Both North and South Pagi present a slightly different picture from
Siberut and Sipora. The Pagi islands rise more steeply from the sea,
so much so that the ten fathom curve runs very close along the shore.
There are a few villages principally along the east coast but the popu-
lation is not large. Apparently the islands are principally covered with
original forest. North Pagi lies about twelve miles south of Sipora.
There are very few inhabitants and little is known of the islands.
Except for the collection made by Dr. Abbott and Mr. Kloss in 1902
there has been only the short trip of Mr. Menden in 1937. It is unfor-
tunate that no field notes were made on these two visits, for undoubt-
edly the Pagi Islands are extremely interesting from the faunistic and
geological point of view.

Twelve miles southeast of South Pagi there is a small coconut-cov-
ered island called Sanding which apparently is little more than a wide
exposed reef. It has never been visited by a scientist. South of Sanding
the two hundred meter curve ends. Some fifty-five miles southeast
of Sipora there is a small pinnacle called Mega, two miles long and

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 317

surrounded by a fringing reef. It is low and densely wooded and rises
from the one thousand meter curve. This island too has never been
scientifically explored.

Enggano, the most isolated island of this archipelago, lies about one
hundred eighty miles south east of Sipora. It is about seventy miles
from the nearest part of the Sumatra coast. The island is twenty miles
long and approximately ten broad. A range of hills runs from north-
west to southeast through the center of the island, expressing thus the
direction of upthrust throughout the group. Enggano lies entirely on
its own shelf, the surrounding seas reaching a depth of three thousand
meters. Enggano is heavily wooded and the interior is so marshy and
difficult to penetrate that communication between the villages is said
to be by boat. It is sparsely inhabited, the 1919 census having shown
only four hundred seven persons on the island. The main trade of the
island is in copra, and the principal settlement is on Aduwe, one of the
three little islands in Enggano Bay. This group is called Pulu Dua.

FAUNAL LIST OF THE BIRDS

In the following list of the birds of the west Sumatra islands, the
arrangement of the families has been that of Wetmore (A Systematic
Classification for the Birds of the World, Smith. Misc. Coll., 99, 7,
1940, 11 pp.), while the arrangement of the species within the families
has been that of Chasen (1935). All measurements are in millimeters,
the wing pressed flat against the ruler. All weights of small birds are
in grams. Where used, wing-tail ratio refers to the percentage of the
length of the tail as compared to that of the wing. Wing tip index
refers to a similar percentage of the length of the shortest primary as
compared to that of the longest primary. Color notes follow Ridgway.

Of the total of seventy-five families found on Sumatra and generally
in the vicinity of these islands, forty-six families are represented from
this archipelago with certainty. Of the missing families some members
of the Phaethontidae, Sulidae, Rostratulidae, Indicatoridae, and Arta-
midae, may be expected to turn up on these islands in the future.
Notable, however, is the lack of representatives of the Megapodiidae
and Phasianidae, which do not occur there and presumably never have
been found on these islands.

318 bulletin: museum of comparative zoology

Family SULIDAE, Boobies, Gannets

The only record for this family is a dubious one which I have not
included in my definite list of species of the islands. There is every
reason to suppose, however, that boobies may occur around these
waters from time to time.

? Sula sula rubripes Gould

Chasen (1935) records this form from the Pagi Islands. I can find
no record of a specimen from this area except that in the British
Museum Catalogue (Vol. XXVI, p. 434, 1898), which lists a juvenal
male from Nassau Island, south of Sumatra, August (Dr. Coppinger),
Voyage of H.M.S. ''Alert". In the report on this voyage (British
Museum, 1884), Dr. Coppinger notes in the summary of the trip (p. 3)
that the boat touched at Nassau Island in the Danger Group northeast
of Samoa, after leaving Tahiti on August eighteenth. There is no
mention of a stop near south Sumatra during the later part of the trip,
although a stop was made at Singapore in November of the following
year. More evidence is needed to prove the existence of this form off
the west Sumatran islands.

Family PHALACROCORACIDAE, Cormorants

The only record for this family is from Nias. There is no indication
as to whether or not the Little Cormorant is a resident in the islands .

1. Phalacrocorax niger (Vieillot)

Listed by Biittikofer (1896) from Nias on the basisof Nieuwenhuisen
and von Rosenberg's report (1863). This species has not since been
recorded from any of the islands.

Family ARDEIDAE, Herons

2. Ardea sumatrana sumatrana Raffles

Recorded from Simalur, Babi, Tuangku, Nias, and the Pagi Islands.

The record for the Pagi Islands is by Chasen and Kloss (1935).
I can find no specimen on which it is based. A nest with two well-
grown young was found by Abbott on Simalur in November. A female
adult in the collection of the National Museum was secured on the
same island, October 23, 1902. Wing, 434.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 319

3. Ardea purpurea manilensis Meyen

Found on Xias, Pini (Batu Islands), and Enggano.

An immature specimen was taken on Enggano by Modigliani, May
19, 1891. An adult female collected by Dr. Abbott, November 5, 1904,
on the same island was labeled "iris yellow". Wing, 351.

4. Butorides striatus spodiogaster Sharpe
Synonym: Butorides striatus sipora Chasen and Kloss

Occurs on Simalur, Nias, Siberut, Sipora and North Pagi.

Three specimens from Simalur, Nias, and Sipora measure: wing
(worn), c? 182.5, 9 170.5, o 180.5; tail, d* 63, 9 63.5, o60; culmen,
J1 63, 9 60, o 60.

Oberholser (1912, p. 1) named two races, actophilus from North
Pagi Island and icastopterus from Simalur, on the basis of larger size
and darker color. The types measure :

actophilus 9 ad., icastopterxis c? ad.,

January 4, 1903. December 10, 1903.
wing 196 191

tail 70 68

culmen 66.5 65.5

Another female from North Pagi, collected December 31, is also
large (wing 190, tail 67.5, culmen 62.5). These birds presumably
represent a northern race found in Siam and China which migrates
during the winter to the southern islands. Chasen and Kloss (1926)
have described sipora as being darker on the neck than javanicus,
although similar in size. The three small specimens agree with this
description, as well as appearing darker on the back and scapulars
than a large series of javanicus. However, they are unfortunately in-
separable from spodiogaster Sharpe (Bull. Brit. Orn. CI., 3, 1894,
p. xvii) of the Andamans and Nicobars. A female from Little Nicobar
collected March 3 has a wing of 180 and agrees well in color with the
west Sumatra islands.

The names carcinophilus and carcinophonus Oberholser (Journ.
Wash. Acad. Sci., 14, 1924, p. 294) and abbotti Oberholser (U. S. Nat.
Mus. Bull. 159, 1932, p. 14) are synonyms of javanicus.

5. Butorides striatus actophilus Oberholser
Synonym: Butorides javanicus icastopterus Oberholser

Migrant from the Asiatic mainland recorded from Simalur (icastop-
terus) and the Pagi islands in December and January.

320 bulletin: museum of comparative zoology

6. Ardeola bacchus (Bonaparte)

Recorded by von Rosenberg from Nias (Biittikofer, 1896) and a
sight record (Richmond, 1903) for Simalur.

7. Egretta garzetta nigripes (Temminck)

There is a record of this species for Nias in Biittikofer 's list based
on Nieuwenhuisen and von Rosenberg (1868).

8. Egretta eulophotes (Swinhoe)

A rare migrant from China where it has been seriously reduced by
plume hunters. Recorded from Sipora by Chasen and Kloss (1926)

9. Demigretta sacra sacra (Gmelin)

Occurs on Simalur, Tuangku, Nias, Pulo Tello, Sipora, the Pagi
islands, and Enggano.

Two specimens collected by Dr. Abbott on Simalur and Pulo Mirbau
off Enggano measure: 9 ad., wing 262, tarsus 68; c? im., wing 266.5,
tarsus 69.5. The immature bird is in the mottled juvenal plumage
for the white phase discussed by Mayr and Amadon (Amer. Mus.
Novit. No. 1144, Oct. 13, 1941). Curiously enough, the wing measure-
ment of the adult female is smaller than any of the series of three
hundred seventy-eight specimens measured by them.

Of the thirty-three known adults collected on these islands, nineteen
are described as being in the grey phase, thirteen in the white phase,
and one (Salvadori, 1892) in the mottled phase.

The reef heron probably nests on the small islets off the coasts of
the larger islands. Junge (1936) records a set of eggs taken July 3
on an islet in Sinabang harbor, Simalur.

10. Mesophoyx intermedia intermedia (Wagler)

The single record for this species is for Nias, Biittikofer (1896)
founded on von Rosenberg's list.

11. Nyticorax nycticorax nycticorax (Linnaeus)

Found on Nias (Biittikofer 1896) and Enggano. Two males and
a female from the latter island measure: wing, c? 270, 287, 9 281.
One male is in an erythristic state of plumage similar to that recorded
for N. n. hoactli by van Rossem (Auk, 53, 1936, p. 322). The cheeks

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 321

and nape are tinted with buff. The upper part of the back is dark
bronzy brown shading posteriorly to "Carob brown". The wings and
tail are greyish buff. The second male approaches this but is paler and
more greenish on the back. The female is nearly normal. The occur-
rence of erythrism in this species indicates that Xycticorax caledonicus
may be a form in which this type of mutation has been established as
a dominant. It will be interesting to observe the apparently new
colony of caledonicus breeding in East Java, (Hoogerwerf, Bull. Raffles
Mus., 12, 1936, p. 120), to see if the birds hybridize with nyciticorax.
The colors of the soft parts are recorded as follows : iris straw yellow;
bill black, base and lores greenish (cT), lores slaty (cf), and base
pinkish fleshy ( 9 ) ; feet pale greenish yellow (d\ 9 ), straw yellow (cf ).

12. GORSACHIUS MELANOLOPHUS MELANOLOPHUS (Raffles)

Two females collected by Dr. Abbott on Enggano seem to be the
only record for this species in the west Sumatra islands. One specimen,
collected November 22, 1904, is in immature plumage. The soft parts
are as labelled: "iris pale greenish yellow; feet brownish green; bill
black above, pale brown beneath."

13. Ixobrychus sinensis sinensis (Gmelin)

An immature male collected by Abbott on Nias, March 28, 1903, is
the second record for that island.

14. Ixobrychus cinnamomeus (Gmelin)

A single male is recorded from Enggano June 22, 1936, by June
(1938).

15. Dupetor flavicollis flavicollis (Latham)
Recorded from Xias by von Rosenberg.

Family CICOXIIDAE, Storks

The first member of this family found on these islands proves to
belong to a widely-distributed form throughout the Malay area.

16. Dissoura episcopus stormi (Blasius)

North Pagi. An adult male and an immature female taken on North
Pagi in December by Menden are in the collection of the Museum of

322 bulletin: museum of comparative zoology

Comparative Zoology. They are the first specimens of this family to
be recorded from the west Sumatra islands. The male specimen has
the following notes on the colors of the soft parts : "iris brown, bill and
feet red" (but basal third of bill is black in the skin). The immature
has the colors noted as: "iris dark brown, feet red, bill red-brown"
(not black basally in skin).

I am indebted to Mr. James L. Peters for the identification and the
information on the labels.

Family ANATIDAE, Ducks and Geese

The Javan Tree Duck is the only species recorded on the west
Sumatra islands so far. It is a wandering species throughout the
Greater Sunda islands. Very likely Dendrocyqna arcuata occurs here
from time to time, while the absence of Anas castanea gibberifrons
from this area is definitely puzzling, (see Ripley, Auk, 59, 1942, p. 98).

17. Dendrocygna javanica javanica (Horsfield)

Biittikofer (1896) lists this as the probable race encountered by
von Rosenberg. When I was on Nias in June, 1939, I was told that
ducks were common on Nias at certain seasons.

Family ACCIPITRIDAE, Hawks, Eagles

Sixteen members of this family have been recorded from these
islands of which ten are presumably residents and six migrants. Five
of these resident forms are presumed to be identical with those found
commonly throughout the Malayan area. Of the remaining five,
Spizaetus cirrhatus vanheurni is described as being smaller than its
nearest relative. The other four forms belong to the very plastic
Spilornis cheela "rassenkreis". The two races from the northern
islands, abbotti and asturinus resemble birds from the Andaman and
Nicobar Islands. The race from the Batu Islands has been united
with the population from south Sumatra. The southernmost race,
that from Sipora and the Pagi islands has characters which markedly
resemble those found in Java, Bawean and Celebes. In all these Spi-
lor?iis forms there are marked color differences.

18. Pernis apivorus ptilorhynchus (Temminck)
Siberut (Chasen and Kloss, 1926, p. 279.).

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 323

19. HALIASTUR INDUS INTERMEDIUS Blyth

Found on Simalur, Babi, Tuangku, Nias, and the Pagi islands.

Junge (1936) notes that this bird was found nesting on Simalur in
May in a coastal grove of casuarinas at a height of 20 m. above the
ground. An immature bird was collected at the spot.

20. Accipiter badius poliopsis (Hume)

An adult female collected on November 20 is recorded from Nias
by Buttikofer (1896).

21. Accipiter soloensis (Horsfield)

Taken by Abbott on Simalur in October and January. Chasen
(1935, p. 71) lists this species from Nias on the basis of Biittikofer's
record of Accipiter poliopsis. From the description of the specimen,
however, it is impossible to be sure that he did not have a specimen of
poliopsis.

22. Accipiter trivirgatus trivirgatus (Temminck)

Recorded for Nias. A male collected March 26 is in extremely worn
plumage showing no sign of new feathers. A female collected on the
same date is also very worn, but there are a few new feathers coming
in on the sides of the neck and nape. Another female collected March
23 is renewing some of the tail feathers and the inner primaries. A
few of the nape feathers and wing coverts are new also. Soft parts:
iris, cf , "bright yellow"; 9 "orange yellow"; feet, d* "yellow", 9
"pale yellow" ; bill, 9 "black above, leaden beneath". The stomach
of the female (March 23) contained lizards.

23. Accipiter virgatus gularis (Temminck and Schlegel)

Simalur, Siumat, Lasia, and Nias. Four immature females from
Simalur and Siumat and Lasia measure: wing, 182, 185, 185.5, 188.
Two of the specimens had been eating small birds. This species has
been collected from November 25 to February 12.

24. Spizaetus nipalensis alboniger (Blyth)

Found on Pulu Asu, off Simalur, Bangkaru (sight), Nias, and North
Pagi. Chasen's record (1926) for S. cirrhatus limnaetus on the Pagi
islands presumably referred to this species. Three immature males

324 bulletin: museum of comparative zoology

from Pulu Asu, Nias, and North Pagi measure: wing, 296, 308, 310.
They were taken December 25, March 13, January 6. Soft parts:
iris, "greyish-yellow, lemon yellow"; feet, "dull lemon yellow"; claws,
"black".

25. Spizaetus cirrhatus vanheurni Junge

This race was described by Junge (1936) on the basis of five speci-
mens from Simalur. The author notes that the birds are much smaller
than S. c. limnaeetus, but failed to compare them with S. nipalensis
alboniger. In measurements and in coloration, these birds seem very
similar to alboniger.

26. Ictinaetus malayensis malayensis (Temminck)

Listed by Biittikofer from Nias on the basis of Nieuwenhuisen and
von Rosenberg (1863). This mountain forest species has never been
encountered since on the west Sumatra islands.

27. Haliaeetus leucogaster (Gmelin)

A bird of the small islands off Simalur; Asu, Djawi djawi, Babi-
ketchil, all in Sinabang bay, Siumat, and Lasia, Babi, Bangkaru,
Tuangku, Nias (vide Chasen, 1935), Sipora, and the Pagi islands.

Two males were collected on Asu and Siumat in December. A nest
was seen by Abbott on Siumat on Christmas Daj^.

28. Spilornis cheela abbotti Richmond

Simalur. This well-marked race was collected by Abbott (1901) and
van Heurn (1913) at all months of the year. The wings of nine males,
six females, and one unsexed, measure: cfcf 330-362, 9 9 319-359,
o328.

Soft parts: "iris yellow; bill horn blue, brownish at tip; cere orange
yellow; inside of mouth leaden; feet dirty yellow."

Stomach contents: snakes, small crab, centipede, lizards.

Apparently an abundant bird on Simalur, this race is very distinc-
tive in appearance. The amount of spotting on the under parts is
variable. In some specimens it extends up onto the breast; in others it
is confined to the abdomen. The marking on the breast and throat is
far more noticeable than in the paler forms, due to the contrast be-
tween the dark blackish brown and reddish brown bars.

No pale individuals have so far been taken. Meise's recent revision
of this genus (J.f.O., 87, 1, 1939, p. 65) indicates that all these popula-
tions should be considered races of cheela.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 325

29. Spilornis cheela asturinus A. B. Meyer

Nias. Three males collected by Abbott measure: wing, 291, 291.5,
299.5 All three birds are in worn plumage with a few new feathers
appearing among the greater wing coverts (one male) and secondaries
(another specimen). Soft parts: iris bright yellow, feet dirty orange
yellow, cere yellow. Stomach contents: snakes and small centipedes.

A pair collected by myself for the Academy of Natural Sciences
weighed, d* 15 oz., 9 1 lb. 4 oz.

This species is common on Nias, where it may be found perching in
a high tree on the edge of an open field or moving slowly over the
gardens or hedgerows in search of its prey.

From minimus of the Northern Nicobars, which it closely re-
sembles, this form differs by the finer barring and spotting of the
lower underparts.

30. Spilornis cheela batu deSchauensee and Ripley

Tello, Tana Massa. Meise (I.e. 1939) in his splendid revision of
the genus Spilornis lumps Malay Peninsula, Sumatra, Java, and Bali
birds all under bassus, giving for wing measurements, 348-394. I am
inclined to believe that the name bido should stand for Java and Bali
birds, which are large (384, 391), and dark, with distinctly blackish
throats. Granted the last character is allelic in these birds, still it
seems to be quite constant in the Java population.

The birds from south Sumatra, on the other hand, are uniformly
small, distinctly paler in tone than bido, although in this last character,
they do not differ from typical bassus.

When de Schauensee and I described batu (1940, p. 401.), we had
only north Sumatran material available for comparison. An adult
from Tana Massa in the National Museum's collection (sex indet.
no. 179622) does not bear out our contention that this is a pale race.
It is as dark in tone as any specimen from Sumatra or the Malay
Peninsula. However, it does agree with the Academy's two birds in
being small. Four males, three females, and one sex indet. from the
east coast province of Sumatra agree with these Batu Island birds in
size and color. They come from the Katenan River, the Siak and
Little Siak rivers, Makapan, and Pulu Padang. Their measurements
are: <?d\ wing 229, 331, 346.5, 352, tail 204, 206, 217 (molt), 220;
9 9 , wing 340, 350, 354, tail 218, 221, 221 ; o wing 346, tail 215.

As can be seen, these measurements are all below 360 for the wing
and 230 for the tail. North Sumatra birds, on the other hand, measure

326 bulletin: museum of comparative zoology

much larger. Six males and four females from Atjeh measure: wing,
c? 360, 365*, 365, 388.5, 390*, 410.5; 9 371*, 375, 380, 388; tail, <? 231
255*; 9 253*. Similarly seven Malay Peninsula birds measure wing,
cf 355 (1, worn) - 397 (374); 9 385^07 (392); & 230-259 (244.6),
9 242-264 (251.6).

Under these circumstances, I prefer to restrict the name bassus to
birds from the Malay Peninsula and Sumatra south to the province
of Oostkust. I propose also to unite birds from Oostkust south with
Batu Island birds as batu, modifying the description of that race
(I.e. 1940,) to read: Pulo Tello and South Sumatra birds "differ by . . .
smaller size".

31. Spilornis cheela sipora Chasen and Kloss

Sipora and the Pagi islands. A single female from Sipora has a
wing measurement of 310 and the soft parts are indicated as: "iris
and facial skin yellow; bill pale grey; feet yellow".

This race is distinctive in having the feathers of the crest largely
white with black tips giving a noticeably piebald effect. The breast
is solidly colored, smoky bronzy black. In its coloration, sipora shows
a close relationship to the black-cheeked strain of Spilornis which con-
tinues southward with bido of Java, baweanus of Bawean, and rufipectus
of Celebes.

Family FALCONIDAE Falcons
Two species have been recorded from the islands, both migrants.

32. Falco peregrinus calidus Latham

A migrant collected on Nias and Enggano and probably seen on
Simalur (Richmond, 1903). A male from Nias collected in March has
a wing of 306.5. A female from Enggano collected in November
measures 376. The latter specimen is molting the greater wing coverts.
The soft parts of the male are given as: "iris dark brown; eyelids
yellowish grey; feet bright yellow".

33. Falco tinnunculus subsp. ?

A winter migrant recorded from Nias in November by Biittikofer

(1896).

• U.S.N.M. birds.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 327

Family RALLIDAE, Rails, Coots, Gallinules

Four species occur on the west Sumatra islands all apparently
identical with wide-ranging Malayan forms.

34. Rallus striatus gularis Horsfield
Synonym: Hypotaenidia striata reliqua Oberholser

Simalur. Two females collected in October and December are
identical with specimens from Java and Singapore. The soft parts are
marked as: "iris pale yellow brown, red; bill pinkish red, tip horn
brown; feet dull brownish purple". Found in the rice paddies.

(Rallus jouyi Stejneger is by no means a synonym of gularis. It is
a giant pale race. The male type and a female from Shanghai measure :
wing, cf 135, 9 134.5; tail, cf 50, 9 49; culmen, d" 43.5, 9 42.5.)

35. Rallina fasciata (Raffles)

Nias, Sipora, and Enggano. A male from Enggano collected in
November measures: wing 128.5. Soft parts: "iris and eyelids red; feet
vermilion."

36. Amaurornis phoenicurus javanica (Horsfield)
Synonym: Amaurornis phoenicura cleptea Oberholser

Found on Simalur, Nias, Tello, Siberut, Sipora, the Pagi islands, and
Enggano.

A common bird of the flooded rice fields and swampy meadows near
streams. Found also in the sago swamps on those islands which have
them. These birds have a harsh rattling call.

Four males from Simalur, three females from Nias, including the
type of cleptea, and four females from Sipora are inseparable on the
basis of size.

37. Porphyrio poliocephalus indicus Horsfield
Recorded from Nias by von Rosenberg (1878).

Family CHARADRIIDAE, Plovers

Four species have been found on these islands of which three are
migrants. The resident form has been separated on the basis of color,
a character which I fail to distinguish.

328 bulletin: museum of comparative zoology

38. Pluvialis dominica fulva (Gmelin)

A common migrant recorded from Simalur, Nias, Batu islands,
Siberut, and Sipora, September through April.

39. Charadrius peronii Schlegel
Synonym: Charadrius peronii chaseni Junge

A resident species recorded from Simalur and Nias. A female from
Simalur (M.C.Z. 178151) was collected in May. Junge described the
race chaseni from Simalur as being "darker on the upper parts, less
rufous on the head especially in the 9 9 . Also the rufous colour
behind the white neckcollar and on the sides of the breast is much more
pronounced than in p. peronii. The bill in chaseni is slightly heavier."

A female from Nias collected in February (two months earlier than
Junge's Simalur series) is a good deal darker than the Simalur female.
It is in much fresher plumage. A molting female from the Philippines
collected in January has new feathers as dark as the Nias bird, while
an October male from Mindanao is considerably darker than any other
bird in our series. As far as the character of the rufous breast band
goes, the two west Sumatra island birds stand midway in a series of
seven females. I cannot find any significant size difference in the bill.
I fail to see, therefore, how chaseni can be upheld.

40. Charadrius mongolus atrifrons Wagler

A migrant recorded from Simalur and Tello; this species probably
occurs on all the small beaches. Two females from Simalur collected
in December measure: wing, 130, 134.5.

41. Charadrius leschenaultii Lesson

A winter migrant. There are records for Simalur, Nias, Pini, Tello,
and Sipora from August through December. A male and two females
collected on Simalur in December measure: wing, cf 140, 9 140.5,
142.5.

Family SCOLOPACIDAE, Snipe, Woodcock, Sandpipers
Twelve species have been found on these islands, all of them migrants.

42. Numenius phaeopus phaeopus (Linnaeus)

A migrant taken on Simalur, Sipora, and Siberut. A female taken
on Simalur in November has the rump pure white. A male from

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 329

Sipora collected in February has a few spots of grey brown among the
white rump feathers. For the time being, I think it is better to call
these birds phacopus. There seems to be a good deal of intergradation
between the two forms.

43. NUMENIUS PHAEOPUS VARIEGATUS (Scopoli)

Reported from August till June on Simalur, Nias, Pini, and Tello.
There is a sight record of one or other of the races of whimbrel for the
Pagi islands.

44. Numenius arquatus orientalis C. L. Brehm

A migrant taken on Tuangku and Nias. A female from Tuangku
(January) measures: wing, 282.

45. Limosa lapponica baueri Xaumann

The single record of this migrant for the west Sumatra islands is
Xias, October (Blasius).

46. Tringa totanus eurhinus (Oberholser)

This winter visitor has been taken on Simalur, Nias, and Tana
Massa from September until May. A female from Nias collected in
March has a wing measurement of 158.

47. Tringa nebularia Gunnerus

A migrant reported from Simalur and Nias in August, September,
and October.

48. Tringa glareola Linnaeus

This winter bird was found on Simalur from January until April.

49. Actitis hypoleucos (Linnaeus)

The common sandpiper of the islands from August until April.
Recorded with certainty only from Simalur, Babi, Nias, Tello, and the
Pagi islands.

50. Arenaria interpres interpres (Linnaeus)

A female collected on Simalur in December has a wing measurement
of 148.5.

330 bulletin: museum of comparative zoology

51. Capella stenura (Bonaparte)

During migration recorded from Simalur, Nias, Sipora, and (sight
record) the Pagi islands. Two males and three females from Simalur
and Nias measure :wing, & 135, 138; 9 134.5, 135, 135.5. These birds
were collected during December, February, and March.

The Pintail Snipe is a common bird in these islands during the
winter, being found wherever there are recently drained rice fields or
along the edges of streams, sometimes in the short grass bordering
roads built through swampy areas.

52. Erolia ruficollis (Pallas)

A casual winter visitor, found on Simalur and Nias in December
and October.

53. Erolia testacea (Pallas)
Blasius (1901) records this migrant as taken on Nias in October.

Family BURHINIDAE, Thick-knees
A single resident species occurs throughout the greater Sunda area.

54. Orthorhamphus magnirostris (Vieillot)

Simalur, Babi, Tuangku, and Nias. A male from Babi collected in
January has a wing measurement of 291.5. Weight, 2.5 pounds. Soft
parts: "iris yellow, tarsi pale yellowish slaty, thighs yellow, claws
black, leaden at base". The Stone-plover feeds on crabs and is common
along sandy beaches and on exposed reefs at all seasons.

Family GLAREOLIDAE, Pratincoles
A single species of this family has been recorded twice as a migrant.

55. Glareola maldivarum Forster
A migrant from northern Asia recorded from Nias and Sipora.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 331

Family LARIDAE, Gulls, Terns

Five species have been noted as occurring on these islands. At least
one species is known to breed here, but there has been no observable
speciation in the case of this family.

56. Chlidonias hybrida javanica (Horsfield)

There is a record for Nias in December (Blasius, 1901), and von
Rosenberg (1878) includes this in his list of species met with from time
to time among the islands.

57. Sterna dougallii bangsi Mathews
Collected on Siumat in May.

58. Sterna sumatrana sumatrana Raffles

Simalur, Pandjang, Djawi-djawi, Siumat, Nias, and Enggano. Eggs
have been taken in May on Simalur and Djawi-djawi. An immature
male was taken off Nias by Abbott, October 30.

59. Thalasseus bergii cristatus (Stephens)

Mentioned by von Rosenberg (1878) as occurring in the seas about
these islands.

60. Anous stolidus pileatus (Scopoli)

Listed by von Rosenberg as seen from time to time among the
islands.

Family COLUMBIDAE, Pigeons, Doves

Twenty-one forms are recorded from the west Sumatra islands. Of
these seven belong to wide-spread species, common throughout the
Greater Sunda area. Thirteen of the fourteen other forms are endemic.
One is considered identical with Sumatran and Malayan subspecies.
Of the thirteen endemic races eleven differ markedly from their cor-
responding relative on Sumatra by larger size and rather different
coloring. Two forms do not differ in size. One, Treron fulvicollis
melopogenys, is poorly characterized and may not be a good race. The
other, Ducula aenea consobrina, differs from D. aenea only in color.

332

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The eleven well characterized forms may be tabulated as follows:

D. aenea M . ruficeps M. phasianella

T. curvirostra

ggano*

zzm

Mentawi
& Paei.

Xias

Simalur

BBZ2Z
i I i i

Sumatra

1. 2. 3. 4. 5. 1. 2. 3. 1. 2. 3. 4. 5. 1. -2. 3. 1. 2. 3. 4. 5.

M H difference in size.
V/J///A difference in color.

Fig. 3

In the above chart Sumatra birds are taken as standard. Their in-
dividual variation is taken as less than the arbitrary unit 1, within
which it is considered that local island populations may vary and still
be considered inseparable. The unit 2 is used to express those varia-
tions in size or color, either one of which by itself is still too slight for
separation.

Above this number the units used indicate the relative degree of
differentiation of the two factors, size and color, of the different island
populations.

61. Treron curvirostra haliploa Oberholser

Simalur. The type, an adult male, measures: wing 145.5; tail 92.5;
culmen 16.5. The soft parts are marked: iris orange, orbital skin yel-
lowish green, feet dull purple. Junge (1936) gives the measurements
of four males as : wing 146-151 ; tail 90-94; culmen 16-17.5. This race
differs from curvirostra by larger size. Thirteen males from Peninsular
Siam and Sumatra (harterti is a synonym) measure: wing 132-139;
tail 77-83.5; culmen 16-18.

Junge (I.e.) notes that a single male from Babi is quite yellowish on
the under surface. His description sounds very much like smicra from
farther south (see also Junge, 1938, p. 340.).

A bird of original forest.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 333

62. Treron curvirostra pega Oberholser

Nias. The type, an adult male, measures : wing 144; tail 88; eulmen
16. Four females measure: wing 134.5, 142, 143.5, 143.5; tail 75, 79,
79, 82; eulmen 16.5, 17, 17.5, 18. This form is larger than curvirostra
and differs both from that form and haliploa by being noticeably paler
on the under parts and by having more grey on the flanks. The soft
parts are recorded as: "iris bright ochreous ; orbital skin apple green,
pale bluish green; feet maroon red; bill yellow; cere red." Weight:
cf, 7 oz.

The female differs as the male.

Specimens with enlarged gonads were taken by myself in June.

63. Treron curvirostra smicra Oberholser

Tello, Tana Bala, Siberut, and Sipora.

The type, an immature male taken February 8 on Tana Bala, meas-
ures: wing 138.5; tail 73; eulmen 17. Four males and a female from
Siberut and Sipora measure: wing d" 141, 141.5, 145, 147.5, 9 148.5;
tail d\ 83.5, 84, 90, 9 86; eulmen & 17.5, 17.5, 17.5, 18, 9 18. Soft
parts: "iris yellow; bill pale green; cere olive; feet maroon red, crimson
lake".

This race is very bright yellow on the under surface. The breast
particularly is a rather rich shade of lemon yellow. It is the brightest
member of the species. On the upper surface it differs from its near
relatives by having the nape more distinctly yellowish green less
greyish green.

The female differs as the male.

Evidently this bird prefers a slightly different type of habitat than
either haliploa or pega, which are found on much larger islands. If
the specimen from Babi, one hundred and sixty miles to the north,
recorded by Junge, proves to belong to this race, it would seem to
indicate that smicra not only prefers a different ecological setting but
also suits its behavior to its location. Small island populations of
pigeons seem to be more subject to sporadic wandering movements
than their large island congeners.

64. Treron curvirostra hypothapsina Oberholser

Enggano. The male type, collected November 21, measures: wing
156; tail 92; eulmen 17. The soft parts are recorded as follows: "iris
vellow; eyelids dull green; bill jade green, base dull green; feet purplish
red."

334 bulletin: museum of comparative zoology

Two other males and a female measure: wing c? 149, 150, 9 141;
tail cf 86, 90.5, 9 85.5; culmen c? 18, 18.5, 9 16.

This again is a larger bird than curvirostra and much brighter yellow
on the under surface. It differs from smicra by being slightly duller on
the under parts, lacking especially the bright suffusion of yellow on the
breast found in the latter subspecies. On the upper surface, too, the
nape is more grayish-green, less tinged with yellow. Junge (1938)
records enlarged gonads in males collected in June and July.

65. Treron fulvicollis melopogenys (Oberholser)

Nias. The type and only available specimen is a female collected by
Dr. Abbott, March 18, 1903. It measures: wing 141 ; tail 85.5; culmen

16. Soft parts: "iris pinkish mauve; bill greenish blue; cere dull
crimson, eyelid gray with yellow edge; feet maroon red."

Two females of fulvicollis from Sumatra and Sarawak measure: wing
145, 150. This is a rather variable species particularly in color. Lack-
ing a larger series, however, I suppose that this single rather small
bright-yellow-throated specimen must be recognized.

66. Treron olax (Temminck)

Recorded by von Rosenberg (1878) as being on "all the larger
islands."

67. Treron vernans miza (Oberholser)

Simalur. The type, an adult female, collected November 22, meas-
ures: wing 152.5; tail 95; culmen 15. Soft parts: "feet deep red."
Three males measure: wing 156, 156.5, 158; tail 98, 98.5, 104; culmen

17, 17.

This is purely a size race. There are no color differences compared
to griseicapilla. Junge (1936) describes the plumage of a juvenal bird
collected in July. Eggs were collected in April, June, and July.

68. Treron vernans griseicapilla Schlegel
Synonym: Dendrophassa vernans mesochloa Oberholser
Synonym: Dendrophassa vernans polioptila Oberholser

Nias, Tana Massa, Tana Bala, Siberut, Sipora, North and South
Pagi, Enggano. The type of mesochloa, an adult female taken on Nias
March 18, measures: wing 145; tail 81.5; culmen 16.5. Soft parts are
recorded as : "iris pinkish-yellow; bill gray; cere dull green; feet maroon

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 335

red." A series of males and females from Nias measure: wing c? 141
155, 9 141, 146.

The type of polioptila, an adult female collected on North Pagi
January 1, measures: wing 148; tail S9.5; culmen 14.5. A series of
males and females from the Batu Islands, Sipora, North and South
Pagi and Enggano, measure: wing cf 148-156, 9 150-152.

These birds differ a good deal in size. Nias birds are more nearly the
size of Sumatran and Malay Peninsula birds, while specimens from
the islands slightly to the south, particularly North Pagi, tend to be
larger. In view of the overlapping measurements, however, it seems
to be much wiser to group these birds all under griseicapilla. None
of the local populations seem to be entitled to subspecific recognition,
either on the basis of standard deviation tabulation or of zoogeography.
Only the Simalur birds are apparently significantly larger. It is in-
teresting to note that a large series of griseicapilla from Sumatra and
the Malay Peninsula preserve far more uniformity in their wing
measurements (cf 144-151, 9 142-150) than do the birds from the
western islands. Presumably this is due to less possibility of swamping
of the island populations.

Junge (1938) records enlarged gonads in a male from Enggano taken
in June. This is a common bird in the islands. It is fond of thick
secondary growth in swampy coastal locations. As with all pigeons,
they are very fond of the fruit of various species of Ficus.

69. Leucotreron jambu (Gmelin)

Recorded from Nias in August (Blasius) and from the same island
by Hartert.

70. Ducula aenea consobrina (Salvadori)

Synonym: Carpophaga Vandepolli Biittikofer

Synonym: Muscadivores consobrina babiensis Richmond

Synonym: Muscadivores aeneus mistus Oberholser

Synonym: Muscadivores aeneus vicinus Riley

Simalur, Lasia, Babi, Tuangku, Nias, Tello, Tana Bala, Siberut,
Sipora, North and South Pagi.

The name vandepolli was founded on a discolored Nias bird (Junge,
1935).

The type of babiensis, an adult male from Babi, measures: wing
248; tail 146.5; culmen, 23. Two pairs from Lasia and Babi measure:

336 bulletin: museum of comparative zoology

wing, cf, 245, 247 (246.6); 9 237, 237; tail, <?, 144, 146.5; culmen,
<?, 24, 26.5.

The race mistus from Simalur was described as being decidedly
smaller than Nias birds. The type measures: wing, 231; tail, 135.5;
culmen, 22.5. The wing measurements of a series of males from
Simalur, 225.5-238 (231.1), as against a series from Nias, 230.5-244
(235.8), show far too much overlap to have any significance.

When the wing measurements of the three males of babiensis are
compared with those of twenty-five males from Simalur, Tuangku,
Nias, Tana Bala, Sipora, North and South Pagi, there is only a small
amount of overlap (2.5) :

wing average

adult c? d\ Babi and Lasia 245-248 (246.6)

" other islands 226.5-247.5 (234.6)

However, the range of variation is very great and accordingly I have
tested them by the formula of t used in the comparison of the means
of two small samples, (Simpson and Roe "Quantitative Zoology",
1939, p. 210). Using this I obtained a value for / on the wing measure-
ments of 1.9. This gives a probability of between .1 and .05 showing
that the deviation is not significant. This bears out the evidence of
a male from Babi recorded by Junge (1936) which measured well
within the range of consobrina.

The race vicinus from the Batu and Mentawi islands was founded
on color: "the breast and hind neck washed with much deeper
vinaceous-lilac." The color of these birds is subject to much variation
through dirt or grease on the feathers. When absolutely clean speci-
mens of each island are compared, however, color variations tend to
disappear.

From aenea these birds differ by lacking the pronounced vinaceous
wash on the head, cheeks, and sides of the neck. As a result, there is
less contrast between the gray head and the white margin around the
bill. The under parts generally tend to be more gray, less tinged with
vinaceous, although the latter tint is noticeable in dirty specimens.
The wider tail coverts seem to average darker in consobrina. The
measuring of wings in these birds is sometimes quite difficult, due to
the wing being incompletely pulled back by the skinners. This fact
undoubtedly accounts for some of the variations in measurements.

Molting specimens were collected in November and immature birds
in October, November, and January. Eggs have been taken on Simalur
in May and June. Immature birds are distinguished by a less well de-

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 337

fined division between the gray of the nape and the bronzy green of
the back. Also the marginal under wing coverts are edged with
rufous.

Soft parts of adults are given as: "iris, deep red; bill, pale leaden,
base dark leaden; cere, dull purple; feet, livia purple." Immature
birds are given as: "iris, sepia; bill, gray; cere, faintly brown; eyelids,
gray edged with pale pinkish brown; feet, purplish brown."

These big pigeons are common and conspicuous on most of the
islands, although on Nias they seem to be confined to the higher parts
of the island where some original forest is still found. The birds are
trapped a great deal by the people either as young taken from the
nest or by the use of bird lime (Malay "gutta"). Pigeons are a staple
of the native diet.

71. Ducula aenea oexothorax (Salvadori)

Enggano. Ten males and two females measure: wing, d", 247-254;
9 , 246, 249; tail, d\ 152-156; culmen, d\ 21.5-25.5.

This race differs from aenea by larger size and by lacking the vina-
ceous wash on the head. In most specimens the tail feathers are dis-
tinctly greenish instead of bluish iridescent in color. On the under
surface, however, there is an ill-defined band of strong vinaceous wash
on the breast. The under tail coverts are dull brownish green.

From consobrina this race differs by larger size and a slightly longer
tail proportionate to the length of the wing (62% compared to 57-
59%). The same characters of vinaceous breast band and more green-
ish under tail coverts apply in this case as well as with aenea.

Soft parts: iris, deep crimson; bill, pale leaden, base dark leaden;
cere, dull purple; feet, dull purplish red.

72. Ducula bicolor bicolor (Scopoli)

Simalur, Lasia, Babi, Tuangku, Nias, Tello, Lago, Pini, Sipora, the
Pagi islands, and Enggano. Specimens from Simalur, Babi, Sipora,
and Enggano collected in October, November, December, and Jan-
uary measure: wing, d\ 224-238, 9 , 223.5-231.5. An immature bird
was taken on Sipora in October. The soft parts are recorded as: iris
dark; bill leaden, black at tip; cere greenish; feet blue.

This is a bird primarily of the very small islands and reefs. They
follow the seasons flying about in small flocks in search of trees in
fruit.

338 bulletin: museum of comparative zoology

73. Ducula bicolor badia (Raffles)

Rosenberg (1878) records encountering this bird on Nias, the
Mentawi islands, and Enggano. It has not been met with since that
time.

74. Columba Argentina Bonaparte

Recorded from Simalur, Sipora, and South Pagi. There is a sight
record for Kokos. Males and females from Simalur and South Pagi
measure : wing, d\ 239-249.5; 9 227, 240. The soft parts are given as:
iris, bright red, yellow suffused with red; bill, greenish horny, dull
purple at base, apple green; cere, dull purple; tarsi, purple; toes, pale
purplish fleshy.

This is a wandering seasonal species not common in collections. It
probably occurs from time to time on all the west Sumatra islands.

75. Macropygia ruficeps simalurensis Richmond

Simalur. Two males and two females measure: wing, cf , 152.5
(type), 149; 9 , 144.5, 146; tail, tf, 164.5 (type), 169, 9 , 155, 164.5;
culmen, d\ 14 (type), 14; 9 , 13, 14.

As well as having certain differences in color noted by Junge (1936),
this race is slightly larger in size than sumatrana.

76. Macropygia phasianella hypopercna Oberholser

Simalur. The type, an unsexed bird evidently a female, measures:
wing, 172.5; tail, 177; culmen, 17. This bird is grease stained on the
upper back and rump. With only one immature specimen, it is im-
possible to determine the validity of this race. However, its resem-
blance to the Nias race is striking.

77. Macropygia phasianella modiglianii Salvadori

Nias. A male and two females measure: wing, cf, 184.5, 9, 183,
185; tail, 9, 184, 184.5; culmen, <?, 17.5, 9, 16, 17.

This race differs from emiliana of Sumatra and Java principally by
larger size. There seems to be a slight reduction also in the amount of
the irregular barring on the breast.

78. Macropygia phasianella elassa Oberholser

Siberut, Sipora, and North and South Pagi. The type, an adult
male from North Pagi, collected November 12, measures: wing, 179.5;

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 339

tail, 171 (molt); culmen, 20. The soft parts are marked: "iris, pink
with an inner narrow blue circle."

This race is distinguishable from the available specimens of modi-
glianii. In the male the upper parts are very similar but specimens of
elassa tend to have more distinct pale tips to the feathers of the back,
scapulars, and wing coverts, giving a slightly barred effect. On the
under surface, the iridescent purple sheen of the breast tends to be
confined to a rather distinct band, whereas in modiglianii, the color is
more generally dispersed. The color of the under side of the tail
feathers seems to be paler, also, and lacking in gloss.

In the female, the color differences are more apparent. The neck
and upper back are distinctly barred black on buff, giving an effect
of more contrast. The tail seems to be slightly darker. On the under
surface there is a distinct tendency to a blackish patch on the throat.
In the male, this spot is variable and sometimes absent. In the large
series of females examined, however, it is always 'present. The rest
of the under surface seems to be distinctly darker than in modiglianii.

79. Macropygia phasianella cinnamomea Salvadori

Enggano. This race represents the farthest step in a cline running
from north to south in the islands. The Nias population differs from
emiliana primarily by size. Farther south elassa has more distinct
barring, a condition which resembles that of immature birds. Also
there is a tendency towards reduction in the amount of violet iridescent
gloss spread over the plumage and the appearance of a blackish throat
patch. In cinnamomea this condition is further exaggerated. The iri-
descent gloss has disappeared. The plumage as a whole appears faded
and washed out. The black barring on the back tends to break down.
The primaries are pale rufous on their outer edges as in immature
specimens of the other races. The black throat patch is present as a
definite adult character.

An immature female is uniformly pale rufous brown all over, except
for the upper back which is irregularly shaded with black, and the
tail and inner margins of the primaries which are blackish.

Measurements: wing, & , 205.5, 207, 9, 196, 203; tail, d\ 181.5,
198, 9 , 173, 179; culmen, d\ 22.5, 22.5, 9 , 20, 20.5.

80. Chalcophaps indica indica (Linnaeus)

A wide-spread form recorded from Simalur, Siumat, Lasia (vide
Chasen, 1935), Nias, Tello, Siberut, the Pagi islands, and Enggano.

340 bulletin: museum of comparative zoology

It will probably be found to occur on all the islands with suitable
forest.

On Nias I saw the Green-wing Dove several times sunning and dust
bathing in the middle of the small island roads.

81. Caloenas nicobarica nicobarica (Linnaeus)

Recorded from Kokos, Simalur, Lasia, Babi, Nias, Batu Ids., and
Mirabau, an islet off Enggano.

This is primarily a bird of the small coral islands off shore, but
Nieuwenhuisen and von Rosenberg record it as common in the
Lagoendi Bay region of south Nias, near the present-day Telokdalem.

Family PSITTACIDAE, Parrots

Seven psittacine forms occur on the west Sumatra islands of which
six are endemic. The species Psittacula alexandri has no representa-
tives on Sumatra. The three races found on these islands are closest
to the population of the Andaman Islands. Like that race they differ
from the bird of the mainland primarily by larger size as well as certain
color characters.

The species Psittacula longicauda found on Sumatra as well as the
Andamans and Nicobars, has a single highly specialized representative
on these islands on Enggano. It differs by much larger size and rather
juvenal color characters.

There are two forms of Psittinus cyanurus on these islands. One,
from Simalur is not only much larger than the Sumatran bird, but
has certain striking color differences. The other, from the Mentawi
and Pagi Islands differs from the Sumatra bird only in larger size.

82. Psittacula alexandri cala (Oberholser)

Simalur. The type, an adult male taken October 21, measures:
wing 185, tail 198.5. The rest of the series collected by Abbott in
October and November measure: wing, d\ 173.5 (worn)-177.5, 9,
168.5-177.5; tail, cf, 179.5-196, 9, 170-182.

This race resembles abbotti from the Andaman islands very closely.
The only tenable difference seems to be that the pinkish color of the
breast, particularly in the females, is more pure, less suffused with
lilac. Other differences can apparently be matched in a large series.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 341

From fasciata this race differs by larger size, lack of any trace of
green on the forehead, and the less intense shade of green on the back.

This species is abundant in the west Sumatra islands nocking to-
gether and nesting throughout the year.

83. Psittacula alexandri major (Richmond)

Lasia and Babi. The type, an adult male from Babi, January 14,
measures: wing, 196.5, tail, 230.5. Other birds from the two islands
measure: wing, d\ 189.5, 196.5, 201.5, 9, 186, 188.5; tail, d\ 218.5,
229, 9 , 196, 200.5.

This is a very large race differing from cala primarily in size,
although the males seem to show a reduction in the light bluish suf-
fusion on the lower abdomen and vent. One female (U.S.N.M.
179112) has a thin ring of pink extending from the sides of the neck
dorsally around the nape between the bluish-gray crown and the
green back.

Soft parts: iris, pale yellow with an inner dull green ring; upper
mandible (cf), red; cere, greenish leaden, dull greenish; feet, pale
green, pale dusty green.

84. Psittacula alexandri perionca (Oberholser)

Nias. The type, an adult male collected February 22, measures:
wing, 194.5; tail, 211.5. Another male and two females measure : wing,
tf, 182, 9 , 177, 183.5; tail, <?, 189.5, 9 , (molt), 170.

This race occupies a somewhat intermediate position as regards cala
and major in size. Like major the males have less of the heavy bluish
suffusion on the lower abdomen and vent. The color of this area is
somewhat brighter than in major.

In June these birds were nesting, using holes made high up in the
trunks of dead coconut palms.

85. Psittacula longicauda modesta (Fraser)

Enggano. The two species, alexandri and longicauda, have an extra-
ordinary distribution in the greater Sunda area. It is particularly
interesting that alexandri, a species found away from the mainland,
principally on small islands, should have reached Java. This argues
a long history for the range of the species. And yet it is a representa-
tive of longicauda which has reached Enggano, the most isolated of
the west Sumatra islands.

342

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The race modesta. occupies one end of a cline starting with longicauda
of the mainland, Sumatra, and Borneo. The cline consists principally
of color characters which may be summarized as follows:

Subspecies

Loral
stripe

Crown

Nape

Back vs.
Scapulars

Size
Wing d" cf

longicauda

indicated

emerald
green

pink, like
cheeks

strong
contrast

small

defontainei

indicated

yellowish

emerald

green

paler

strong
contrast

larger

tytleri

present

yellowish
green

lacking

weaker
contrast

larger

nicobarica

present

chromium
green

lacking

weak

contrast

larger

modesta

extending

over

nostrils

indistinctly
green

present

very

pale

weakest
contrast

largest
199.5-209

The series collected by ^Abbott measures: wing, cf, 199.5-209
(205.6), 9, 195.5-209.5 (201.1); tail, d71, 182 (worn)-233, 9, 131-
151.5

Molting specimens were collected in October and November and a
fully grown immature female November 11. The soft parts are marked
as: iris, yellow, inner circle green; upper mandible, red, lower, horn
brown, black (cf); feet, greenish leaden, slaty-green, greenish.

A male and a female are heavily stained on the under parts, pre-
sumably from eating fruit.

86. Psittinus ctanurus abbotti Richmond

Simalur and Siumat. This very distinct form lies somewhere on
the borderline between a species and a subspecies. Primarily it reflects
the speciation trend of these islands by being much larger than cya-
nurus. Also, however, there are important color characters which set
it off from the typical race.

Adult male: the crown is a more uniform blue than in cyanurus,
lacking the gray wash on the nape. Also there is a sprinklingof greenish

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 343

iridescent feathers on the forehead and the ocular area. The grayish
black back and scapulars of cyanurus are lacking, being replaced with
green. The mid portion of the back is bright blue instead of pale dull
purplish blue and the rump and upper tail coverts are green instead of
blue. On the under surface greenish yellow replaces the yellowish gray
breast, abdomen, and vent, and the blue thighs of cyanurus.

Adult female: above this bird is almost uniformly green, lacking
the dull brown head of the typical race. Below, also, abbotti is uni-
formly greenish yellow.

The type, an adult male taken in December, measures: wing, 145;
tail, 58.5; culmen (from cere) 22.5; tarsus, 13. Iris, pale yellow; cere,
dull green; feet, greenish.

A series of males and females measure: wing, c? , 139-147.5 (143.5);
cf, im., 139; 9 , 134.5-141 (137.6).

An immature male collected in October has an indistinct tinge of
blue on the forehead. An adult male (U.S.N.M. 179109) has a spot of
dull red similar to that of the humeral patch among the blue feathers
of the nape.

This is a rather tame little parrot, common locally when the wild
fig trees are fruiting.

87. Psittinus cyanurus pontius Oberholser

Siberut, Sipora, and South Pagi. This race is distinguished from
cyanurus by larger size. In a large series, any color differences are
shown to vary within the species. A male from Siberut (U.S.N.M.
279754) has a very dark blackish back. Soft parts: "iris, pale yellow;
upper mandible, red tipped with yellow, lower, olive horn; cere, dull
leaden olive; feet, olive."

The type, an adult male collected in December, measures: wing,
127; tail, 50.5; culmen, 20.75. The rest of the series measures: wing,
<? , 127-134.5 (131.5); 9 , 129-132; tail, cf , 50.5-54. Birds from Su-
matra and the Malay Peninsula measure: wing, cf , 117-125 (121.6),
9, 115-123; tail, 44-47.

88. Loriculus galgulus galgulus (Linnaeus)
Synonym: Loriculus galgulus lamprochlorus Oberholser
Synonym: Loriculus galgulus dolichopterus Oberholser

Tuangku, Nias, Pini, Tello, Siberut, Sipora, Enggano.

The type of lamprochlorus, an adult male collected on Nias, March

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14, measures: wing, 79; tail, 32.5; culmen (from cere), 10.5. This race
was described by Oberholser (1912) as being "decidedly smaller, and
the colors averaging paler; female with green color more yellowish".
A series of birds from Sumatra, Borneo, and Java (three specimens
which may well be cage birds) measure: wing, cT, 78.5-83 (80.3).
The color characters also disappear in a series.

The type of dolichopterus, a female from Enggano collected Novem-
ber 6, measures: wing, 83.5; tail, 34.5; culmen (from cere), 11.5. A
series of females from Sumatra, Borneo, and Java (see anted) measure:
wing, 77-84.5 (81.4). This race was described as being larger and
darker, but these differences do not appear in comparison with a
large series of birds. Junge (1938) reached the same conclusion.

The wings of some of these specimens have not been fully pulled
back after skinning. In the case of the type of dolichopterus this may
have accounted for the impression that it was a larger bird.

Family CUCULIDAE, Cuckoos

Fifteen species have been recorded from these islands of which two
are migrants. Of the remaining species four are endemic on the islands.
Of these four races, two differ by color alone while the other two differ
only by larger size. All the races are closely related to representatives
of the species found on Sumatra.

89. Cuculus fugax fugax Horsfield

Pini, Siberut. An immature specimen from Siberut collected
October first, apparently belongs to this race. Chasen's record for
nisicolor (1935) may well be founded on this and another immature
bird taken by Kloss. The bird measures: wing, 167.5; bill (from gape),
26.5. Some of the slaty adult feathers are beginning to appear on the
forehead, cheeks, and nape. There are several white feathers also on
the nape.

90. Cuculus micropterus concretus S. Muller

Tana Bala. An adult male collected February 11, tail lacking, is
the first record of this species from the west Sumatra islands. It meas-
ures: wing, 191.5; culmen, 22.5. The soft parts are recorded as: "iris,
ochreous, rim of eyelid yellow; lower mandible, gray; gape, yellow;
feet, yellow." Compared with a male from Trang and a female frojn
east Sumatra collected in January and February, this specimen is dis-

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 345

tinctly more gray on the back, wing coverts, rump, and upper tail
coverts. The underparts seem to be identical. It is unfortunate that
no more specimens were secured.

91. CUCULUS SATURATUS SATURATUS Blyth

A migrant from southern Asia discussed by Junge (1931) in a careful
revision of the species. He lists two specimens of this race from
Simalur taken in February. The other record for the west Sumatra
islands is Nias (Biittikofer 1896, p. 171).

92. Penthoceryx sonneratii fasciolatus (S. M tiller)

Pini, Tana Massa. A male and a female collected by Kannegieter
in September and October on these two islands measure: wing, d*
(worn) 114, 9 111. These birds are indistinguishable from two males
from Trang, a locality certainly within the range of malayanus Chasen
and Kloss.

93. Cacomantis merulinus threnodes Cabanis and Heine
Synonym: Cacomantis merulinus subpallidus Oberholser

Nias, Siberut, and Enggano. The type of subpallidus, an immature
male from Nias, is a rather pale bird, but an adult male from Nias is
similar to a male from Borneo. I believe that this slight difference in
coloration is due to individual variation. Two adult males from Nias
and Siberut measure: wing, 101, 101.5.

The characteristic ascending notes of the brain-fever bird are often
heard on Nias in open areas of garden and cultivation.

94. Cacomantis variolosus sepulcralis (S. Mtiller)

Simalur, Enggano. A male molting into adult plumage and an im-
mature female were taken by Abbott on Simalur, December 14 and
November 30. The soft parts were noted as:' "iris, d* gray brown; 9
reddish brown becoming gray externally; eyelids, d1 greenish yellow;
bill black, brownish yellow at base of lower mandible; inside of mouth
orange; feet yellow."

95. Chalcites xanthorhynchus xanthorhynchus (Horsfield)
Only recorded from Simalur and Sipora.

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96. SURNICULUS LUGUBRIS DICRUROIDES (Hodgson)

A female collected on Nias March 15 evidently belongs to this large
raigatory race. It measures : wing 141.5, tail 135, culmen 22. The soft
parts are labelled: "iris dark brown, bill black, feet dark leaden."

97. Surniculus lugubris barussarum Oberholser

Tana Bala. The type of barussarum, an adult female taken on Tana
Bala in February, measures: wing 136, tail 127, culmen 22. Another
female taken in the same month measures: wing 137, tail (worn) 120,
culmen 22. Both birds have distinctly forked tails, the lateral feathers
in the type being 6mm. longer than the central ones.

This race was named (1912, p. 5.) as being smaller with the bill
relatively larger than I. lugubris. Actually two males of lugxibris from
east Java have the following measurements: wing 124, 127, culmen
20.5, 21. Robinson (Birds Malay Penin. /, 1927, p. 140.) gives meas-
urements for brachyrus of the Malay Peninsula, Borneo, and Sumatra
which are very close to these two birds from Tana Bala. A female
from Singkep Island, east Sumatra taken in May measures only:
wing 128, tail 123, culmen 20.5. If this specimen may be taken as an
example of typical brachyurus, then I presume that banissarum repre-
sents a large island form. Larger series may show, however, that these
measurements are not distinctive, in which c&sebarussarum must stand
as the name for the birds from the southern Malay Peninsula, Borrfeo,
Sumatra, and the Batu Ids.

98. EUDYNAMIS SCOLOPACEA SIMALURENSIS Junge

Kokos, Simalur, Lasia, Babi. Junge (1936) characterizes this race
as differing from malayana by, "smaller wing and in the 9 also by
the darker coloured and bigger spots on the upper side and the more
strongly red-brown washed under parts".

As to size I cannot entirely agree with this diagnosis. A male from
Babi measures: wing 211, tail 197, culmen 36, while a male from
Simalur in the collection of the Museum of Comparative Zoology has
a wing measurement of only 195 according to Mr. Peters. A series of
males of malayana measure: wing 203-212.5. However, a single pos-
sibly immature female from Simalur is distinctly more rufous above
and below than any female of malayana. In this respect it is very
close to females of mindanensis. Fortunately the bill of mindanensis

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 347

is noticeably smaller. The record for Lasia is a sight one, (Richmond,
1903).

99. Eudynamis scolopacea malayana Cabanis and Heine

Nias, Tello, Tana Massa, Pagi islands. Two males, an immature
male and three females from Tello and Tana Massa measure: wing,
c? 217.5, 218; 9 im. 213, 9 209, 212, 213.

There is a good deal of individual variation in size, bill dimensions,
and color in this species. These birds are large but not significantly
so. The soft parts are recorded as: iris red; bill gray, greenish gray;
feet gray.

The record for the Pagi islands is not founded on any specimen so
far as I know, but undoubtedly these birds could be found on almost
any of the small coral islands of the area, especially in the breeding
season.

100. Rhopodytes sumatranus rodolphi Ripley

Pini. The type, an adult male collected October-November 1896,
measures : wing 154, tail 237, culmen 37.5. The soft parts are recorded
as: iris light blue, bill yellowish green, feet gray.

Although only a single specimen has been taken on the west Sumatra
islands, its bill measurements are significantly larger than a series of
specimens from Sumatra or the Malay Peninsula and adjacent islands.
In color there seems to be no perceptible differences between this and
Sumatra or southern Malay Peninsula birds. This specimen has very
little rufous on the abdomen, but I believe that individual variation
or skinning will account for it.

101. Rhinortha chlorophaea facta Ripley

Tana Massa. The type, an adult male collected February 20, 1903,
measures: wing 123.25, tail 182, culmen 29, tarsus 28.5. A female
(A.N.S.P. 56256) collected August 24, 1896, measures: wing 124, tail
179.5, culmen 30.5, tarsus 28.5. The soft parts are given as: "iris, 9
blue gray; bill, cf dark greengage green, 9 , green; feet, c? slaty, 9
gray."

These birds are noticeably larger than any other specimens of the
species in the Museum's series. Except for Banguey I. off the coast
of North Borneo this bird has never been reported on any of the small
islands in the Greater Sunda area.

348 bulletin: museum of comparative zoology

102. Rhamphococcyx curvirostris oenicaudus
(Verreaux and Verreaux)

Siberut, Sipora, North and South Pagi. This race differs from
erythrognathus of the Malay Peninsula and Sumatra by having the
greenish iridescence on the upper surface rather darker and more
bronzy, extending up onto the nape and crown and serving to darken
the gray of those areas. The tail lacks the terminal brown patch
characteristic of the other members of the species.

Below the greenish iridescence extends up over the abdomen. The
under surface of the tail is greenish blue not brown. There seems to
be no size difference within the species.

Five males, seven females, and three sex unidentified, measure:
wing, cf , 166-181, 9 165-178, o 171-179. Soft parts are recorded as:
"iris, c? bluish white, pale blue, pale china blue, 9 dark; bill, maxilla
apple green; nostril black, pale green; mandible green, base dull
crimson, dark red; orbital skin scarlet, red; feet plumbeous."

103. Centropus sinensis bubutus Horsfield

Nias, Siberut. Four males and one unsexed from Nias collected in
February and March measure: wing, & 212, 215, 218, 221, o 220, tail
<? 272, 278, 278, o 285. The variation in these birds from purplish
to greenish iridescence about the head, neck, and breast varies with
the freshness of the plumage.

Family TYTONIDAE, Barn Owls

The only species of this family found in the area is the small Bay
Owl of the greater Sunda Islands which is not uncommon on Nias.

104. Phodilus badius badius (Horsfield)
Nias. This species has only been recorded from Nias where it is
not uncommon. For further comments and field notes see de Schau-
ensee and Ripley (1939, p. 402.).

Family STRIGIDAE, Owls

Six forms of owls are found on the west Sumatra islands, all of which
are apparently endemic. Two differ from their congeners in Sumatra
and Borneo, primarily by color differences. Three of the remaining

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 349

forms are distinctly larger than their nearest relatives on the Malay
Peninsula or Sumatra, as well as differing slightly in color. One race
is smaller than its Sumatran congener.

105. Otus scops umbra (Richmond)

Simalur. The type and unique specimen, an adult male, was col-
lected by Abbott November 29, 1901. It measures: wing 143, tail 61,
culmen 20, tarsus 24. The soft parts are recorded as: iris greenish
yellow; bill pale brown, black at tip; feet pale fleshy brown. The wing
formula is 4>3>5>2>6>7>1.

This bird is presumably in the rufescent phase. The outer scapulars
have conspicuous white spots and the feathers of the lower abdomen,
vent, and flanks are freckled and mottled with white, russet, and black
in a way that seems characteristic of the scops group.

The stomach contained insects.

106. Otus scops enganensis Riley

Enggano. A single female, the type, was collected by Abbott, No-
vember 12, 1904. Its measurements are: wing (worn), tail 73, culmen
23, tarsus 30. The tips of the primaries are completely worn off, a
condition that was not mentioned in the original description. The
wings of five birds collected by deJong (Junge 1938) measure: c? ,
160, 163; 9 , 163, 163, 166. The soft parts in the type are recorded as :
"iris greenish yellow, feet pale brownish fleshy."

The type is a bird in the rufescent phase of plumage. In his dis-
cussion of this form, Junge (I.e.) showed that the brownish or gray-
brownish individuals are very close to malayanus and sunia. The
under parts, however, are more uniformly dark rufous than any
rufescent individual of scops I have seen, except for the type of umbra,
which is the most uniformly dark of all.

107. Otus bakkamoena mentawi Chasen and Kloss

Siberut, Sipora, North Pagi. In their original description (1926),
Chasen and Kloss neglected to state that this race differed from true
lempiji by much larger size. A series of these birds measure: wing,
c? 157> 165, 9 165, 165.5, 9 im. 160; tail, tf 77.5, 82, 9 72, 80, 80,
9 im. 76. Ten specimens of lempiji from Java, Sumatra, and the
Malay Peninsula measure: wing, cf 141-150.5 (145.2), 9 140-151.5
(145.1); tail, d* 65-70 (67.2), 9 65-67.5 (66.3)..

350 bulletin: museum of comparative zoology

The soft parts of these birds are listed as: "iris, brown (3), yellow
(2), feet gray." The color differences noted in the original description
are very noticeable in this series.

108. Ketupa ketupu minor Biittikofer

Nias. Compared with ketupu from adjacent areas, three males and
a female from Nias show the following measurements:

wing tail

Nias cf 304-330.5 (316.1) 143-164 (153.5)

9 304 147

Sumatra1 cf 320-349 (338.6) 158-171 (165.4)

9 335, 338 158

Borneo c? 312-343 (330.1) 153-164.5 (158.4)

9 — —

Java & 341-357 (348.4) 157-169 (164)

9 358, 362 170, 176

From the above it will be seen that minor is a tenable race but that
there is a good deal of overlap with Sumatra and Borneo birds. Two
of the Nias males are darker on the breast than any specimen of our
series of ketupu.

Three young were taken in late March. The plumage is rather uni-
form dark buffy all over except for the thighs which are whitish. Both
upper and under parts are marked with the narrow dark brown shaft
streaks found in the adult only on the under parts.

109. Strix leptogrammica nyctiphasma Oberholser

Bangkaru. The type, an adult male collected in January, measures:
wing 299.5, tail 154. A female taken at the same time measures: wing
307, tail 167.

The type differs from a male from the Kapuas River, west Borneo
(w. 298, t. 168) by having the cheeks somewhat paler and the dark
barring on the underparts slightly heavier, the bars being closer to-
gether. In size there is no difference and the color differences are so
slight that a larger series may well show these forms to be identical.

From myrtha of Sumatra these birds differ primarily by smaller
size. This bird will probably be found one day on Babi and Simalur,
provided there is some original forest left for them.

1 have padded out my series with measurements taken in the same manner by Mayr (Bull.
Raffles Mus., No 14, 1938, p. 13) and de Schauensee and Ripley (Proc. Acad. Sci. Philadelphia.
91, 1939, p. 324.)

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 351

110. Strix leptogrammica niasensis Salvadori

Xias. A male and female (M.C.Z. 194794, 194795) collected in
July 1937 by Barbara Lawrence measure: wing, a71 273, 9 280.5;
tail, d" 151.5, 9 156.5. The male is an immature probably completing
the post juvenal molt {vide Robinson, Birds Malay Benin., 2, 1928,
p. 36). The feathers of the posterior part of the crown and nape and
a few feathers in a ring on the neck below are pale whitish buff in color
and fluffy.

The iris is recorded as brown by Biittikofer, chestnut by Salvadori
(1887). Salvadori (I.e.) lists the bill and feet as pearl gray or sky blue.

Family CAPRIMULGIDAE, Nightjars

There are records of three forms from these islands of which one
has been separated on color characters, and one, Eurostopodus tem-
minckii, shows an unconfirmed tendency towards larger size. The
endemic form, Eurostopodus viaCrotis jacobsoni has no close geograph-
ical relatives except cerviniceps from the Malay Feninsula.

111. Eurostopodus temminckii (Gould)

Nias. Two males taken by Abbott in March measure: wing 211,
213; tail 133.5, 135. Two males from Johore and Banka measure:
wing 201, 203; tail 117, 126, while three males from north Sumatra
are recorded by de Schauensee and myself (I.e., 1939) as having
wings of 197, 198.5, and 201.5 mm. However, Robinson and Kloss
(Journ. Fed. Malay States Mus., 11, 1924, p. 242) give measurements
for males up to 218.

In these two specimens the buffy whitish subterminal bars forming a
ring on the nape are narrower than in any other specimens in the
National Museum series. Also one specimen (180824) has very large
black spots on the crown.

112. Eurostopodus macrotis jacobsoni (Junge)

Simalur. According to the description (1936) a darker bird than
macrotis from the Bhilippines or cerviniceps of the Malay Beninsula.
It is most interesting to find this population so isolated from the rest
of the species. Further collecting might reveal that it occurs in the
Andamans or Nicobars.

352 bulletin: museum of comparative zoology

113. Caprimulgus affinis affinis Horsfield

Recorded from Nias by Nieuwenhuisen and von Rosenberg (1863)
under the name C. maculatus.

Family MICROPODIDAE, Swifts .

Six races of swifts are found on the west Sumatra islands of which
two are endemic. One race differs in color alone, and the second differs
in color and smaller size. Both races are closely related to a wide-
spread form found throughout the greater Sunda area.

114. Collocalia lowi lowi (Sharpe)

Simalur, Babi (sight), Nias. A male from Simalur taken May 5 is
listed by Junge with a wing measurement of 131.

115. Collocalia fuciphaga fuciphaga (Thunberg)

Nias. A single female taken in June is the only record of this species
for the west Sumatra islands.

116. Collocalia vestita vestita (Lesson)
Synonym: Collocalia fuciphaga aerophila Oberholser

Simalur, Nias, Sipora. The type of aerophila, an adult male from
Nias, collected March 16, measures: wing 115; tail, shortest rectrix
48, longest rectrix 53. A male and a female from Simalur and a female
from Sipora measure: wing, d71 112.5, 9 116.5, 116; tail, <? 46 and
50, 9 41.5 and 47, 49 (incomplete). Compared with a male from
Johore, I can see no significant differences in these specimens.

117. Collocalia esculenta cyanoptila Oberholser

Simalur. Junge (1936) records three specimens taken on Simalur
in April and July as belonging to this race. He gives their measure-
ments as follows : wing, cf 107, 109, 9 102; tail, cf 41, 41. The type
and another female of cyanoptila from Bunguran-L, Natunas, measure:
wing, 9 103.5, 103.5; tail, 41.5, 39.5 (molt). Both birds are in rather
worn plumage which accounts for the admixture of purple and greenish
gloss mentioned by Oberholser in his original description (Proc. Acad.
Nat. Sci. Phila., 58, 1906, p. 205).

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 353

118. Collocalia esculenta vanderbilti de Schauensee and Ripley

Nias. The male type and a female measure : wing, cf 99, 9 97; tail,
cf 39, 9 37. This race is smaller than oberholseri and somewhat bluer
above. These little swiftlets are common over jungle trails and roads,
especially in the early morning and evening.

119. Collocalia esculenta oberholseri Stresemann

Tana Massa, Sipora and North Pagi. The type, an adult male from
North Pagi, taken in November, measures: wing 105.5, tail 42.5.
Other birds from Sipora and North Pagi measure: wing, cf 105;
9 98, 105; o 105, 105.5; tail, cf 42.5; 9 40.5, 41.5; o 42, 42.5.

From cyanoptila this race differs by being more greenish above and
with the breast and upper abdomen paler below with more tips among
the brownish gray of that area.

Family HEMIPROCNIDAE, Tree Swifts

Two races of Tree Swifts are found on these islands of which one is
endemic. It differs from its large island and mainland relatives by
larger size.

120. Hemiprocne longipennis perlonga (Richmond)
Synonym: Hemiprocne longipennis ocyptera Oberholser
Synonym: Hemiprocne longipennis thoa Oberholser

Simalur, Nias, Pini, Tana Massa, South Pagi, Enggano. The type
of perlonga, an adult female, was collected on Simalur January 2,
1902. It measures: wing 183.5, tail 112.5. The type of ocyptera, an
adult male collected March 23, 1905 on Nias measures: wing 169.
The type of thoa, an adult male from Pini taken March 7, 1903,
measures: wing 177.5, tail 102. These last two races were described in
a lucid fashion by Oberholser (1912, pp. 7, 8) without being compared
to each other, although they came from closely neighboring islands.

A total of seventeen specimens from the west Sumatra islands have
wing measurements as follows: cf 164-180 (173.1), 9 169.5-183.5
(174.9). Of this series the birds from Nias are the smallest: cf 164-
176 (169), 9 169.5. This is probably due to the fact that Nias of all
the islands of the group seems to be the most closely connected to
Sumatra from a faunistic point of view. Whether in the case of Hemi-

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procne this might be due to swamping is certainly a possibility in the
case of this distinctly aerial species.

I have examined forty-one specimens of longipennis from the Malay
Peninsula, Rhio and Lingga islands, Sumatra, Natunas, Anambas,
Borneo, Banka, and Java. Seventeen males measure: 159-170.5
(164.3). Exclusive of molting birds, twenty-three females measure:
155-173.5 (166). From these figures it will be seen that perlonga is a
tenable subspecies. However, it seems to me that in general this
species has been split into far too many subspecies. The type of
anochroa Oberholser, an adult female from the Natunas, has a greenish
suffusion in the chest region and in general is somewhat darker than
average on the underparts. But in this it can be matched by birds
from the Malay Peninsula, Sumatra, and Borneo.

Stresemann's original description of the race harterti (Novit. Zool.,
20, 1913, p. 339) indicates that he did not have any Bornean material,
for he gives no measurements of specimens from there. Individuals
from Borneo show every variation between the two extremes of colora-
tion of the underparts of this species. Some birds are paler gray on
the throat and breast with more white on the lower abdomen. Others
are darker gray with less white on the abdomen. Northern Malay
Peninsula birds seem to be the darkest, while Java birds are the
palest. In between them is every variation of what is, after all, a
rather slight range of variation. There is no difference in size. Under
these circumstances it seems to me that it would be wiser either to
recognize several microscopic degrees of variation and name all the
small local populations, or else, and this I think preferable, to lump all
these birds as longipennis. This would leave the species arranged as
follows :

Hemiprocne longipennis coronata (Tickell)
India, Ceylon, Siam to Indo-China. A distinctive rufous-throated
race.

Hemiprocne longi pennis longipennis (Rafinesque)
Peninsular Siam to Sumatra, Borneo, Java, and Bali. Dark to
paler gray below.

Hemiprocne longipennis perlonga (Richmond)
West Sumatra islands. A large race.

Hemiprocne longipennis wallacii (Gould)
Celebes to the Sula islands. More bluish on the upperparts.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 355

121. Hemiprocne comata comata (Temminck)
Synonym: Hemiprocne comata stresemanni Neumann

Nias, Pini, Tello, Tana Massa, Tana Bala, Sibe'rut, North Pagi. A
series from these islands measures: wing, cf 126-130 (127.6), 9 123
(worn)-134 (128.5). I fail to note any of the differences mentioned
by Neumann.

Family TROGONIDAE, Trogons

Two trogons occur on the west Sumatra islands. One from Nias is
an endemic race differing slightly from its nearest relative on Sumatra
and the Malay Peninsula in color and a tendency to have a larger bill.

122. Harpactes duvaucelii (Temminck)

The only record for this species is Tana Massa (de Schauensee 1940).
A male and a female taken by Kannegieter measure: wing, c? 108,
9 105.

123. Harpactes oreskios nias de Schauensee and Ripley

Nias. In the original description (1940, p. 404.) this race was said
to differ from uniformis by lacking the orange wash on the chest of the
male and having narrower white bars on the wing coverts and second-
aries. Three males and a female from Nias, when compared with birds
from Peninsular Siam and the Langkawi islands, do not show these
characters. All three of the males have a rich orange suffusion indicat-
ing that the type may not be fully adult. However, these Nias birds
do have somewhat darker more brownish crowns than the birds with
which they have been compared. Mr. de Schauensee writes me (in
litt.) that this is also true of the type. As well as this, these birds
show a tendency to a slightly larger bill. The culmen of the four Nias
specimens measures: c? 17, 17.5, 18.5; 9 18. A series of uniformis
measures: c? 15, 15, 16, 16, 17; 9 14.5, 15, 16, 16, 17.

This trogon is found in heavy secondary jungle at low heights above
the ground, often in substage growth. It has a very distinctive bark-
ing call.

Family ALCEDINIDAE, Kingfishers

The west Sumatra islands have records of twelve species and sub-
species of kingfishers. Of these two are possibly migrants leaving ten
certain resident forms. There are five distinguishable endemic forms
three of which differ primarily by size and two of which differ in color

356 bulletin: museum of comparative zoology

from their nearest relatives. Four of these birds are most closely re-
lated to Sumatran subspecies, while one, Ceyx erithacus captus, is
almost indistinguishable from C. e. motleyi of Borneo.

Ramphalcyon capensis

This is an interesting and widespread species ranging from Siam
down through the Greater Sunda islands to some of the Lesser Sundas.
Oberholser (Proc. U. S. Nat. Mus., 35, 1909, p. 657) has revised the
genus and ended by describing several races of capensis. The addi-
tion of further specimens to the National Museum's collection has
shown, I think, that this action was unwise, as the variation in plumage
of these birds is considerable.

At least in the Malay Peninsula and island part of its range, fresh
plumage in this species is assumed in July, August, or early September.
At this time the color of the pileum may approach sepia to bistre.
The feathers of the upper back, scapulars and wing coverts are rather
bright greenish blue at this time. After eight or ten months, as the
feather tips become frayed and the color of the pileum bleached, very
noticeable changes are brought about in the color of the upper parts.
March to June or July birds have the pileum colored from whitish-
buff or pale fawn to ochraceous-buff. The color of the upper back,
scapulars and wing coverts changes also, this time not due to bleach-
ing, for there is no pigment change as tests under sodium, helium, and
ultra-violet lights have shown. Instead the frayed and sun-heated
feather seem to have become physically changed, contracting the
diffraction grating with resulting effects similar to those observed in
certain beetles. At this time the color of these parts varies from dull
cerulean to indigo to purplish blue.

Birds from the Malay Peninsula and the northern part of Sumatra
tend to be somewhat darker buff on the under surface. Perhaps they
should be kept under malaccensis. Borneo, Billiton, and Java birds
tending to be rather pale beneath, could, it seems to me, quite well
be lumped together as capensis. There is no constant difference in
size in a series of measurements of more than thirty specimens.

From these birds the west Sumatra island populations differ in
being somewhat paler and more washed out on the upper surface and
by a tendency to larger size. The blue which is so bright in capensis
and malaccensis varies in these specimens from dull brownish China-
blue in a September female from Siberut, to pale indigo in a March
male from Nias.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 357

124. Ramphalcyon capensis simalurensis (Richmond)

Simalur. The type, an adult male collected by Abbott November
29, measures: wing 144.5, tail 92.5, bill (from naris) 07. 5. Two other
males and three females measure: wing, cf 143.5 (molt), 143.5 (143.8);
9 151, 155.5; tail, 9 96, 99; bill, d1 65.5, 68.5; 9 70.5, 71.2.

Stomach contents are noted as: "small fish, prawns, and crabs."
This is a common bird of the mangroves and small creeks near the sea.

Compared to malacccnsis, this series is equally dark buff below, but
on the upper surface it is very different, lacking the bright blue or
greenish blue of the upper back, scapulars, and wing coverts.

Compared to birds from the other west Sumatra islands, these
Simalur birds appear uniformly dark buff below in the same way that
malaccensis differs from capensis. The upper parts also appear notice-
ably duller and browner than the majority of other west Sumatra
island birds, but this condition may be in part due to the fact that all
the Simalur birds were collected between October 23 and November 30.

125. Ramphalcyon capensis sodalis (Richmond)

Synonym: Ramphalcyon capensis nesoeca Oberholser
Synonym: Ramphalcyon capensis isoptera Oberholser

Babi (sight), Tuangku, Nias, Pini, Tello, Siberut,Sipora, North and
South Pagi. The type of sodalis, an adult female collected January
25, measures: wing 162, tail 100.5, bill (from naris) 73.5. Another
adult female from Tuangku measures: wing 160, tail 101.5, bill 73.2.
This race was named on the basis of size, but females later added to
the Museum's collection from other islands have equalled these
measurements.

The type of nesoeca from Nias is an adult male collected March 15.
It measures : wing 148.5, tail 93, bill (from naris) 68.5. The range of
this race was given (1909, p. 674) as Nias and the Batu islands, and
the characters were: "pileum paler, back, wings, and tail much brighter
and more bluish." In four of the eight specimens this is true, but as
these are March birds taken at a time when the pale pileum and
brighter blue is characteristic of the state of wear of the plumage, I
do not feel 50% is a good enough average difference for separation.
However, it is certainly worth noting that in the slightly brighter
color of the upper parts these four Nias specimens indicate a closer
affinity with Sumatra than is the case on the other islands.

An adult male taken December 30 in Sikakap Strait between North

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and South Pagi is the type of isoptera. It measures: wing 153.5, tail
94, bill (from naris) 68.5. This race was described (1909, p. 671) as
being larger than simalurensis with the under parts lighter, the lower
back and rump more greenish. In size these birds agree with the birds
from the other islands, as well as in having paler underparts than
simalurensis. I can find no constant difference in the greenish char-
acter of the rump.

Measurements of a series of sodalis by localities follows :

place

wing

tail

bill (from naris)

Tuangku 9

160, 162

100.5, 101.5

73.2, 73.5

Nias c?

146-152.5 (148.7)

89-93 (91)

68.2-72.5 (70.1)

163.5

101

68.5

Batu cf

155

—

92.5

Siberut 9

156.5

97.5

Sipora cf

150.5-158.5 (154.1)

95.5-101 (98)

66.5-70.5 (69)

Pagi c?

150-153.5 (151.5)

89-96.5 (93)

63-70 (67.5)

153, 161

91.75, 95

68, 71.5

126. Alcedo atthis bengalensis Gmelin

Simalur, Tuangku (sight), Nias, Pini, Siberut. Two pairs from Sim-
alur and Nias measure: wing, cf 70.5, 69.5; 9 70, 70. These birds
were taken in November, February, and March.

127. Alcedo meninting meninting Horsfield
Synonym: Alcedo meninting subviridis Oberholser
Synonym: Alcedo meninting callima Oberholser
Synonym : Alcedo meninting proxima Richmond

Simalur, Tuangku, Nias, Pini, Tana Massa, Tana Bala, Sipora,
North and South Pagi, Enggano. The type of subviridis is a male
from Nias collected March 23. It measures : wing 62, bill (from naris)
34. This race was described by Oberholser (1912) as smaller than
meninting with more greenish upper parts. Other birds from Nias
measure: wing, tf1 64.5, 9 63.5, 65.5; bill, cf 34, 9 31.5, 32.5. Birds
from Sumatra measure: wing, <? 61-65, 9 62-66.5; bill, c? 35-36.2,
9 31.5-33.5. Of the four specimens from Nias, only one is especially
greenish on the back and rump.

The type of callima is a male collected on Tana Bala February 8,
measuring: wing 69, bill (from naris) 36.2. It was described (1912) as
larger than meninting with upper parts slightly more greenish. A male

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 359

from Pini and a female from Tana Bala measure : wing, d" 65, 9 68.5 ;
bill, cf 36.5, 9 34.5. These measurements are large, but Junge (1936)
gives measurements for Simalur birds: wing, d* 63, 66, 69; 9 65, 67,
which are largely overlapping. Again the greenish character tends to
disappear with a large series.

The type of proxima is an adult, sex undetermined, from North
Pagi, collected January 4. It measures: wing 69.5, bill 37.5. Other
birds from North and South Pagi measure: wing, cf 67.5-69, 9 71.5;
bill, cf 35.5-37.5, 9 33.5. This race was described as being very
greenish on the back and rump. Two of the specimens are particularly
so — one slightly, and the fourth, the type, can be matched by Sumatra
birds. I think this character is entirely too variable to be a serviceable
criterion for subspecies. These birds are also large, but no more so
than Batu or Simalur birds. Recently Junge (1938) has recorded a
single male from Enggano. The wing measured 63.

From these data, I would be inclined to think that probably these
kingfishers at some time or other reach all the islands of the group.
Also these birds show a tendency to larger size, particularly on Sim-
alur, the Batus, and the Pagi islands. Finally there is a tendency which
I do not consider very significant so far, for some of these birds to have
the iridescent back and rump feathers paler and more greenish.

128. Ceyx erithacus captus Ripley

Nias. This, the only representative of the species from the west
Sumatra islands, is almost identical with motleyi of Borneo. From
this race it differs by a longer bill, slightly larger size, and by the re-
duction of the forehead spot which may be lacking in some specimens.

A not uncommon bird in thick secondary growth or substage vege-
tation usually near small streams. The specimens I saw were all
perched about six feet above the ground.

129. Ceyx rufidorsus jungei Ripley

Simalur, Bangkaru (sight), Tana Massa, Tana Bala. These birds
differ from typical ruflidorsus by larger size. A series measures : wing,
cf 62-64.5 (63.2), 9 62.5-63.5 (63); tail, <? 25-26 (25.5), 9 25.5-26.5
(26); bill, cf and 9 30.5-32 (31.3).

130. Ceyx rufidorsus rufidorsus Strickland

Siberut, Sipora. A male and female adult and an immature male,
taken October 28, are in the collection. They measure: wing, c? 61,

360 bulletin: museum of comparative zoology

9 62; tail, d* 25, 9 26; bill (from naris), tf 25, 9 26. These birds
are certainly close to jungei in size, but Chasen and Kloss' measure-
ments (1926) for Siberut and Sipora birds are so small that, without
seeing the rest of their series, I would rather leave them for the time
being as rufidorsus. The fact that one of the specimens in the National
Museum from their collection is a subadult male inclines me to be-
lieve, however, that some of their other measurements may refer to
immature specimens. There is a sight record for Ceyx from the Pagi
islands, which may refer to this species.

131. Halcyon coromanda minor (Temminck and Schlegel)
Synonym: Entomothera coromanda pagana Oberholser

Simalur, Lasia, Nias, Tana Massa, Siberut, North Pagi. The type
of pagana, an adult male collected January 4 on North Pagi measures :
wing 110, tail 62, bill (from naris) 44.5. Three females from Siberut
and North Pagi measure: wing 107.5, 109.5, 110.5. The soft parts are
listed as: "iris dark, sepia; bill and feet coral red, coral, blood red."
These birds are indistinguishable from minor as is also the type of
neophora, a bird from Tapanuli Bay, northwest Sumatra.

132. Halcyon chloris laubmanniana Grote

Synonym: Halcyon chloris cyanescens * (Oberholser)

Synonym: Sauropatis chloris chloroptera Oberholser

Synonym: Sauropatis chloris amphiryta Oberholser

Simalur, Babi, Nias, Pini, Lago, Tello, Tana Massa, Siberut. Speci-
mens from this area measure as follows (t stands for type) :

wing

bill (from naris)

Simalur o"

115, 116.5, 117.5 (t),

117.5

41, 43, 44.5 (t), 44.5 (2)

117

Nias c?

113.5 (t)

42.5 (t)

114.5

120

43.5

North Pagi

111-116.5 (113)

41-43.5 (41.8)

The type of cyanescens from east Sumatra, an unsexed bird collected
July 28, measures: wing 113, bill 43.5. Specimens from Sumatra,
Banka, and Java measure up to 115.5 (wing) and 43.5 (bill). The
original description of chloroptera from Simalur (Proc. U. S. Nat. Mus.,
55, 1919, p. 379) compared it to cyanescens, but said the birds had

1 Not Halcyon cyanescens Cabanis and Reichenow, 1877.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 361

longer wings and that the upperparts were more greenish, the pileum
was darker, the blaek nuchal hand narrower, the auriculars less green
washed, and the sides and flanks noticeably tinged with huff. In
series these color characters do not appear, nor are the size measure-
ments significant when the amount of overlap is considered.

In the original description of amphiryta from Xias (I.e., p. 382) this
bird seems to stand in an intermediate position between chloroptera
and lavbmanniana. It is smaller than the former and larger than the
latter. In color it is brighter than the former and duller than the
latter. This is a form of microdissection of characters which escapes
me. The plumage changes of these birds are now well enough known
to make comparisons of shades of greenish or bluish color untrust-
worthy. Especially is this difficult when a series of birds, for example
those from Simalur (Oct., Nov., Dec.) or Nias (Feb., March), are
taken all at about the same time. Thus they may present a condition
.of uniformity of plumage which is not really valid for the population
as a whole.

Undoubtedly these birds will eventually be shown to exist on all
the small islands and reefs of this group. They are birds of the man-
groves and beaches and usually omnipresent in these locations. Junge
(1936) lists nestlings from Simalur collected in April and eggs taken
in April and May. Nesting occurs in steep banks along streams, in
hollow trees, and even in termite nests.

133. Halcyon chloris azela (Oberholser)

Enggano and Pulu Dua. This race resembles birds from the Malay
Peninsula and outlying islands, the Andamans,and northeast Sumatra,
in size, but on the average tends to be somewhat darker on the pileum
and more dull greenish on the upper back and scapulars. In view of
the isolation of this island population, it seems strange that these birds
are so slightly distinct. Actually azela appears to be simply a small
race of cyanescens, like that race somewhat duller and darker in color
than the small Malay Peninsula birds.

The type of azela, an adult male collected November 19, measures:
wing 103, bill (from naris) 38.5. A series measures : wing, c? 102 (worn)
-104.5, 9 102-104.5; bill, c? 36-40.5, 9 38.5-41.5.

The races arvistrongi, humii, and davisoni are so far as I can see,
indistinguishable. Sharpe (Cat. Birds Brit. Mus., 17, 1892, p. 277)
describes armstrongi as having green ear coverts and the black band
around the nape obsolete. This description does not seem to be valid.

362 bulletin: museum of comparative zoology

All of the National Museum's series from Peninsular Siam, the Malay
Peninsula and islands, the Andamans, and Belawan Deli, Sumatra,
have at least partially black ear coverts and a well indicated nape
band. Robinson (Birds of the Malay Penin., 1, 1927, p. 99) felt that
the kingfishers from Lower Tenasserim and the extreme north of the
Malay Peninsula were more dull than birds from elsewhere and so
might be called armstrongi. The National Museum has both dull and
bright birds from all localities, which inclines me to believe that they
should all be listed under armstrongi, the name with page priority.

134. Halcyon pileata (Boddaert)

Simalur, Babi (sight), Pini, Siberut, Sipora, North and South Pagi.
This species has not been found breeding south of Bangkok and may
well be a migrant. Specimens taken in November, December, and
January measure: wing, cf 130-131.5, 9 126.5-134. Soft parts were
recorded as: "iris dark brown, bill red washed with brown, feet coral
red." The stomach of one bird contained a lizard.

135. Halcyon concreta concreta (Temminck)

Siberut. The single record for this species is an immature male
taken by Kloss. The wing measurement (1926) is recorded as 112,
and the soft parts: iris dark, bill yellow, culmen black, feet greenish
yellow.

Family MEROPIDAE, Bee-eaters

Two species have been recorded, one of which is only a sight record.
One species which is a migrant has been collected during the winter
months.

136. Merops leschenaulti leschenaulti Vieillot

Dr. Abbott reported seeing this species on Simalur (Richmond,
1903).

138. Merops superciliosus javanicus Horsfield

A migrant taken on Simalur, Tuangku, Nias, Pini, Tello, Tana
Massa, Tana Bala, Siberut and South Pagi. The occurence in these
islands is from September 19, the earliest date, to March 16. Some of
the fall birds are immature. Their plumage is worn, the cinnamon
throat patch is undeveloped, and the upper parts are sprinkled with
blue tipped feathers. Stomach contents, winged ants.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 363

Family CORACIIDAE, Rollers

Two forms, both migrants, occur among the islands in winter. A
third resident form has been described from Simalur. It differs from
orientalis of the Malay Peninsula and Sumatra primarily in color.

139. EURYSTOMUS ORIENTALIS ABUNDUS1 Ripley

Simalur. A male collected by Abbott in December belongs to this
race. The primaries and secondaries are washed with purplish blue
nearly to the end of the outer webs. The specimen measures: wing
194, tail 99, wing-tail ratio 51, wing tip index 36. The soft parts are
recorded as, "iris dark brown, feet red".

Records from Tello, Sipora, North and South Pagi may refer either
to this or the following race, both of which migrate into this area.

140. EURYSTOMUS ORIENTALIS DEIGNANI Ripley

Nias. A male collected by Abbott in March presumably belongs
to this race, although in worn plumage it is not so easy to distinguish
the color characters which separate it from abundus. Certainly there
is no purplish-blue wash on the primaries and secondaries and the
crown is rather blackish. Measurements: wing 1ST, tail 93 (worn),
wing-tail ratio 50, wing tip index 40.

141. EURYSTOMUS ORIENTALIS OBERHOLSERI Junge

Simalur. A single female collected in October, agrees well with
Junge's original description. It measures, wing 187.5, tail 103, wing-
tail ratio 54, wing-tip index 30. Junge (1936) notes that females were
showing nesting activities in January and March. Stomach contents :
large hard coleoptera.

Family BUCEROTIDAE, Hornbills

Two species occur in these islands, one of which is considered to be
identical with Sumatra birds although showing an incipient tendency
to larger size. The other form is endemic, differing from the Sumatra
and Borneo population by definitely larger size.

1 replaces Euryslomus o. catonyx auctorum

364 bulletin: museum of comparative zoology

142. Anthracoceros coronatus convexus (Temminck)
Synonym: Hydrocissa convexa. barussensis Oberholser

Nias, Tello, Tana Massa, Tana Bala, Siberut, Sipora, North and
South Pagi. The type of barussensis is an unsexed bird, obviously a
male with the large wing measurement of 326 and a tail measuring 289.
Birds from the other islands measure as follows :

wing

tail

Nias d" d1

295-319.5 (306.7)

269-294 (282.5)

275

—

Batu Ids. d1 d*

314, 321

285

O (=6)

326

289

9 subad.

279

—

Sipora cT

290

—

Pagi Ids. d* &

282-299 (289.6)

273-282 (277.6)

0(= 9)

263

246

Birds from the Batu Islands are indeed larger than average but I
feel that there is too much overlap to make this race recognizable.
Measurements of over 300 for the wings of Sumatra birds are not
uncommon, and as can be seen from the above measurements there is
a large degree of overlap between Nias and Batu birds. However,these
west Sumatra Island birds do demonstrate in an incipient form a
speciation response by increased size which seems to be so character-
istic of these island bird populations.

Soft parts; iris brown, pale vandyke brown; orbital skin bluish
white; bill ivory, patch on casque and base of bill black; feet dull
plumbeous.

Weight: & 2 lbs., 9 1 lb., 15 oz.

This is a rather shy bird and wherever lumbering or gardening tends
to reduce the original forest, they become much reduced in numbers.
In the south part of Nias where there is little original forest left, they
are now quite scarce.

It is perhaps worth noting that Hydrocissa convexa zamelaena Ober-
holser of the North Natuna Ids. based like barussensis on larger size
appears to be a synonym of convexus. The type measures: wing 312;
tail 308. This seems to be another case of a small island population
tending towards larger size than a mainland or large island population.

143. Crannorhinus leucocephalus megistus Oberholser

Tana Massa, Tana Bala. The type, an adult male collected February
1 1 , on Tana Bala measures : wing 433, tail 283. Another male collected

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 3()5

in the same month has a wing of 408, while a female from Tana Massa
measures; wing 381, tail 251. I have compared these measurements
with ten males and three females from Pulu Payong, East Sumatra,
and west and southwest Borneo. They measure: wing cT 352-405
(385.2), 9 347, 347, 354. The largest Borneo bird measures 401.5.
Comparing these measurements by use of the formula of "t" for small
samples (Simpson and Roe, Quantitative Zoology, 1939, p. 211) I
arrived at a figure of 3.07 for the measurements of the males and 8+
for the females. Thus the probability for the samples being from the
same population is about 01.3% showing that the difference is signifi-
cant. More specimens from Sumatra may show that larger birds occur
there, but for the present I feel that megistus should be recognized.

Soft parts are recorded as follows: cf feet blackish green, soles
dirty yellow; cf im. iris brownish yellow, feet deep dull green, bill pale
blood red at base shading into dull greenish yellow at tip; 9 bill gen-
erally yellow, pale toward tip, mottled at base with dull brownish
green. Skin of throat deep dull slaty blue with a few pale blotches,
orbital skin paler than throat, feet grayish olive.

Weight, c? 3.5 lbs.

I have put this race in the species leucocephalus as I consider Ma-
layan, Philippine, and Celebes birds conspecific. Malayan males differ
by having the nape black instead of rufous chestnut but in two birds
of the National Museum's series this color is not pure black. A number
of small brown feathers are interspersed along the margins of the nape
patch. The shape of the bill is exactly similar although the color is
somewhat different. As leucocephalus is the oldest name I should list
the following well marked races :

Crannorhinus leucocephalus corrugatus (Temminck)
Malay Peninsula, Sumatra, Borneo.

C. I. megistus Oberholser
Batu Ids.

C. I. waldeni Sharpe
Guimaras, Negros, Panay.

C. I. leucocephalus (Vieillot)
Camiguin I., Mindanao.

C. I. cassidix (Temminck)
Celebes.

366 bulletin: museum of comparative zoology

Family CAPITONIDAE, Barbets

Two endemic forms are confined to the west Sumatra islands.
Both differ from their closest relatives on Sumatra primarily in having
larger bills with the addition of certain color characters.

144. Chotorea mystacophanes ampala Oberholser

Tana Bala, Tana Massa. The type, an adult male collected February
11, 1903 by Dr. Abbott measures: wing 104, tail 59.5, culmen 38.5.
Two males and two females measure: wing cf 99.5, 102.5, 9 102,
105.5; tail & 52, 60, 9 55.5, 56.5; culmen & 37, 38.5, 9 37.5, 39.5.

The female with the smaller measurements, collected February 28,
is not fully adult as it lacks the blue tinted throat of maturity. Other-
wise, except for a slight reduction in size of the red patch on the
crown, the plumage is complete and must be presumed to be that
assumed after the post ju venal molt.

Soft parts : feet olive.

This race is chiefly distinguished from typical mystacophanes by the
large bill. Both males and females also seem to have a somewhat
larger red crown patch, and in the female the forehead is more tinted
with blue than in the females from Sumatra and the Malay Peninsula.
Birds from Peninsular Siam seem to show a slight decrease in size
indicating that there is a continuous cline here culminating in the large
Batu Island birds.

In spite of Chasen and Kloss' remarks (Treubia, 14, 1932, p. 14.)
I cannot see the validity of the race humei from Borneo

145. Cyanops australis gigantorhina (Oberholser)

Nias. The type, an adult male collected March 26, 1903 measures:
wing 74.5, tail 38.5, culmen 20.5. A series of four males and three
females measure: wing c? 73.5-81 (76.5), 9 74.5, 75, 80.5; tail cf
38.5-41 (40), 9 37, 38.5, 42.5; culmen & 19-20.5 (20.1), 9 20.5, 21,
21.5. Soft parts are marked as: bill black with gray base, black; skin
of throat dull black; feet olive, green.

This race is characterized principally by the size of the bill which is
noticeably larger than in Sumatra birds, four males and three females
of which measure: culmen & 17-17.5 (17.2) 9 18-18.5 (18.3). The
color of the underparts is slightly variable but tends in Nias birds to
be a purer green especially in the males.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 367

Family PICIDAE, Woodpeckers

Eleven forms occur on these islands of which nine are considered to
be endemic. All nine are related to forms found on Sumatra. Of them
six differ primarily by color characters and three by size. Two of the
size races are larger than their congeners, one is smaller. Of the two
forms considered conspecific with their Sumatran relatives, one,
Dryobates hardwickii subsp. shows a slight inclination towards larger
size measurements.

146. Picus puniceus soligae de Schauensee and Ripley

Xias. This race differs from observandus of the Malay Peninsula,
Sumatra, and Borneo by the greatly reduced amount of yellow on the
prolonged occipital crest feathers. In the type and two specimens of
the series in the National Museum, yellow is quite absent. In the five
other specimens in the National Museum's series, yellow is present,
but it is overlaid by the scarlet vermilion feathers of the crown. Three
males and four females measure: wing d71 126, 126.5, 130, 9 126, 128,
130.5, 132.5.

Soft parts : iris deep crimson, red orbital skin pale blue, slaty blue,
leaden blue; bill black above, ochraceous beneath, lower ochre yellow,
yellow; feet dirty yellow, claws horn brown. Weight 3.4 oz.

As Mayr points out (Bull. Raffles Mus. No. 14, 1938, p. 30.) birds
from Borneo average somewhat smaller than Sumatra or Malay
Peninsula birds. Two females and an immature female from Banka
are larger on the average than typical observandus. They measure:
wing 9 132.5, 136.5, 9 im. 138.

This is a bird of rather thick jungle, often found in dense scrub
growth rather close to the ground.

147. Callolophus miniaceus niasense (Buttikofer)

Nias. A male, three females and one unsexed measure : wing o" 124,
9 123, 123.5, 124.5, o 125.5. Birds from the Malay Peninsula and
Sumatra measure: wing tf1 126, 127, 130, 9 126, 126, 129, 130.

As well as being slightly smaller, this race seems to be more brightly
red on the crown and scapulars.

148. Dryobates hardwickii subspec.

Nias. A single male collected March 23, measures: wing 75, tail
33.5, culmen 17. The latter measurement is larger than a series of

368 bulletin: museum of comparative zoology

birds from Java, Sumatra and Borneo which measure: culmen cf
12.5-16 (14.6) 9 13-16.5 (15.0).

A solitary specimen like this would ordinarily have no significance
in itself, but in the case of a west Sumatra island bird, this slightly
larger bill does point I think to a potential speciation effect in this
species. Further specimens may show such to be the case. Perhaps
this is the form recorded by Nieuwenhuisen and Rosenberg as Picus
percussus (1868).

149. Meiglyptes tristis micropterus Hesse
Synonym: Meiglyptes grammithorax microterus Oberholser

Nias. A series of five males and one female from Nias measure:
wing 90.5-94.5, 9 89. Two males and three females from Sumatra
measure: wing cf 93, 98, 9 91, 92.5, 95. Oberholser 's original brief
description (1912, p. 6) of microterus was "smaller". Use of formulas
(Simpson and Roe, I.e. 1939) shows that the probability of these
specimens being drawn from the same sample is greater than .1%
Thus these small differences lack any significance. In color and mark-
ing these birds are identical.

150. Meiglyptes tukki calceuticus Oberholser

Tuangku. This race was described by Oberholser (1912, p. 6) as,
"like Meiglyptes tukki tukki, but much larger". The type and only
specimen, an adult female collected January 23, 1902, measures: wing
112.2, tail 70, culmen 25. Aside from size there are no other significant
differences.

151. Meiglyptes tukki infuscatus Salvador!
Synonym : Meiglyptes tukki hylodromus Oberholser

Nias. The type of hylodromus an adult male taken March 10, from
Nias measures: wing 97.5, culmen 22. It was described without ref-
erence to Salvadori's race. The rest of the series, five males and four
females measure: wing c? 95.5-103 (98.7), 9 91.5-100.5 (97.3),
culmen c? 22-24, 9 21-21.5. These measurements agree well with
typical tukki.

This race differs from tukki of Sumatra by having less well defined
pale bars on the back and scapulars and by having the barring on the
under parts more reduced on the average.

A male of tukki from Billiton I., southeast of Sumatra (U.S.N.M.

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 369

180492) seems to be an immature bird. Like the young of Dryobates
the feathers of the crown are tipped with a reddish color, in this case
scarlet. Another mark of immaturity may be the pale color of the
abdomen which lacks the characteristic dark rusty wash of the other
specimens.

152. Meiglyptes tukki batu de Schauensee and Ripley

Pini, Tana Massa. A single male in the National Museum's col-
lection has a wing measurement of 109. It differs from the type of
calceuticus by having the barring on the throat more broad and coarse,
and by having the chest more blackish. The series measured by de
Schauensee (1940) seems to indicate that batu is intermediate in size
between tukki and calceuticus.

153. Micro,pternus brachyurus celaenephis Oberholser

Nias. The type is an adult female taken March 29, 1902 measuring:
wing 109.5, culmen 21.5. Six females of badius from Sumatra, Banka,
and Billiton measure: wing 110-122 (113.9).

As may be seen from the above figures, this race does not differ from
badius by being "somewhat larger", (1912, p. 6). It does differ, how-
ever, by being "darker", although its distinctive characters are as
follows : crown with distinct dark centers to the feathers, upper parts
with heavier darker bars; under parts, breast with brownish black
subterminal spots, stomach, flanks, vent and under tail coverts
heavily barred with dark brownish black.

154. Dryocopus javensis parvus (Richmond)

Simalur. The type, an adult male collected December 3, 1901 by
Dr. Abbott, measures: wing 185, tail 124, culmen 38.5. A series of
six males and five females measure: wing d1 172-185 (177.7), 9 172-
182 (177.2).

Soft parts; iris greenish yellow, straw yellow; bill black; feet leaden.

This race is a complete miniature of typical javensis.

155. Dryocopus javensis buttikoferi (Richmond)

Nias. The type, an adult male taken March 18, 1903 measures:
wing 232.5, tail 167, culmen 55. An adult and an immature female
both collected March 25 measure respectively; wing 241, 215, culmen
53, 49.5.

370 ' bulletin: museum of comparative zoology

Soft parts: cf iris bright yellow, upper mandible black, lower
leaden, feet leaden; im. 9 iris straw yellow, feet dusty leaden.

The only discernible difference between this race and javensis is the
lack of the blackish bars found on the thighs of the typical race.

I doubt if these birds are common on Nias at the present time. The
original forest on the island is not plentiful anywhere, although there
is more of it at the north end of the island.

156. Sasia abnormis magnirostris Hartert

Nias. Two males and a female measure: wing cf 52.5, 53, 9 54;
culmen cf 14, 14.5, 9 15. A male from east Sumatra has a wing of
54 and a culmen measuring 13.5, and eight males from Borneo and
the Malay Peyinsula measure: culmen 12-13.5 (13.1). Hartert (1901)
notes that his birds have deeper bills at the base but I cannot find this
difference. Computing the probability of these culmen measurements
by use of the formula of "t" (Simpson and Roe, I.e., 1939) I arrive at
a figure for "t" of 2.7 which is not particularly significant being more
than .02%. However, as there is no overlap, I believe a larger series
would uphold the significance of these measurements especially as the
measurements of four females from Borneo and the Malay Peninsula
are 13-13.5 (13.4), giving when computed by the "t" formula, a prob-
ability for their being identical of only a very little more than .01%.

Soft parts: iris red; ocular area purple, lilac, lilac-purple; bill, upper
mandible worn brown, lower yellow; feet orange, reddish ochre.

Weight: cf 8.5, 11 grams, 9 11 gr.

In contrast to what Robinson says (Birds Malay Penin., 2, 1928,
p. 115.) I found this bird on Nias, not in deep forest, but in secondary
scrub near cuttings and gardens. In June a male with enlarged testes
was collected.

Family EURYLAIMIDAE, Broadbills

Four forms belonging to two species have been recorded from the
west Sumatra islands. Two of the forms, one from each species, are
endemic, differing by larger size, while the remaining forms are
identical with Sumatra birds.

157. Calyptomena viridis viridis Raffles

Nias, Tana Massa. This species has been taken by Modigliani and
Kannegieter. However, it has not been met with since their time. A

RIPLEY: BIRD FAUNA OF THE WEST SUMATRA ISLANDS 371

single male from Tana Massa recorded by de Sehauensee (1940)
apparently does not differ from the typical form.

158. Calyptomena viridis siberu Chasen and Kloss

Siberut, North and South Pagi. A series measures as follows: wing
c? 107.5, cT im. 110, 111, 112 (2), 9 108.5, 112.5, 113. Typical viridis
measures: & 92.5-100.5, 9 98-106.

In size this race agrees with continentis from Peninsular Siam.
However, the single adult male is darker than any specimen of that
form. The females also differ from females of continentis by being
somewhat darker and duller, particularly on the upper parts.

159. Eurylaimus ochromalus mecistus Oberholser

Tuangku. The type, an adult female collected January 29, 1902,
measures: wing 88, culmen 19. Two other females measure: wing
87.5, 87.5, culmen 18.5, 19. These birds are significantly larger than
any specimens of ochromalus measured by me. Apparently they
represent an isolated population on Tuangku, an island lying not more
than twenty-five miles off the west Sumatra coast.

160. Eurylaimus ochromalus ochromalus Raffles

Pini, Tana Massa. Specimens measured by de Sehauensee (1940)
fall within the range of ochromalus rather than mecistus.

wing

culmen

Batu Ids. (f

79.5-83 (81.3)

16.5-18.

79.5-84.5 (81.2)

16-20.

Malay Penin., a1

76-84.5 (80.7)

16.5-20.

Sumatra, Borneo 9

74.5-81.5 (78)

16.5-18.5

There seems to be no difference in color between these specimens.

Family PITTIDAE, Pittas

Two pittas occur on these islands, both of which are presumably
migrants rather than residents.

161. Pitta brachyura cyanoptera Temminck
Synonym: Pitta moluccensis lepta Oberholser

Tuangku, Nias, Sipora. The type of lepta is a male adult collected
on Nias, March 15, 1903 .It measures: wing 121, culmen 25.5. A

372 bulletin: museum of comparative zoology

male and female from Nias and Tuangku taken in January and March
measure: wing c? 121, 9 118, culmen cf 26.5, 9 25. The race lepta
was described as being smaller than moluccensis, "especially the bill".
A series of males from Sumatra measure: wing 120.5-132.5, culmen
26.5-29.

Aside from the difference in size being insignificant, Robinson's
theory ("Fasciculi Malayensis," Zool., 3, 1906, p. 96.) that this is
a migratory form rather tends to mitigate against any attempt to
delimit isolated populations.

162. Pitta sordida cucullata Hartlaub

Nias. Another migrant which has been recorded only by
Buttikofer (1896).

Family HIRUNDINIDAE, Swallows

Two species occur on th

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